L. Poorter, F. Bongers, F.N. Kouamý, W.D. Hawthorne Biodiversity of West African Forests 2004

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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. Kouam, W.D. Hawthorne
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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. Kouam, W.D. Hawthorne
p. cm
Includes bibliographical references and index.
ISBN 0-85199-734-1 (alk.paper)
1. Forest plantsAfrica, West.
2. Plant diversityAfrica, West.
3. Plant diversity conservationAfrica, West.
4. Forest conservationAfrica, West. I. Poorter, L. (Lourens)
QK393.B56 2004
333.9530966dc22
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. Kouam
W.D. Hawthorne
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Table of
Contents
Preface and acknowledgements
Forest
1
2
3
4
5
6
West African forests: introduction
What explains the distribution of rare and endemic West African plants?
Implications for conservation and management
L. Poorter, F. Bongers and R.H.M.J. Lemmens
Forest cover changes in Cte dIvoire 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
15
33
41
F. Bongers, L. Poorter and W.D. Hawthorne
Floristic diversity of closed forests in Cte dIvoire

F. N. Kouam, 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. Belign, W.D. Hawthorne, F. N. Kouam, M.P.E. Parren and D. Traor
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 Cte dIvoire 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 grer et
conserver les forts dAfrique Occidentale (de la Guine au
Ghana).
The project started in 1996 and was jointly prepared by
Renaat van Rompaey, Frans Breteler (Wageningen
University), Dossahua Traor (University of Cocody,
Abidjan) and Vincent Belign (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 Cte dIvoire Vincent Belign and Dossahua Traor
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 Ak-Assi,
Aman Kadio, Vincent Belign, 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 Traor,
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 Genve
(Genve, Switzerland), Musum National dHistoire
Naturelle (Paris, France), Herbier National de Cte dIvoire,
University of Cocody (Abidjan, Cte dIvoire), 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 Genve database), Carel
Jongkind (Combretaceae), Marc Pignal (part of the Paris
database), Roger Polhill (Loranthaceae), Bonaventure Sonk
(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),
Franois Kouam, 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.
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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, Franois Kouam,
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, Franois Kouam,

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, Franois Kouam, 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.
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Forest
Chapters 1-8
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Page 5
L. Poorter, F. Bongers and R.H.M.J. Lemmens
West African forests:

introduction
10:23 AM
H A P T E R
11/11/03
Upper Guinean forests
The rainforests of West Africa have been earmarked as

one of the worlds 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 Cte dIvoire.
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 Rgion guinenne, later renamed
by Monod (1957) as Rgion 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 libro-ivoren of
Aubrville, 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, Cte
dIvoire, 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
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Chapter 1. West African forests: introduction
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, Cte dIvoire, and Ghana. This rainforest belt
is up to 350 km wide, but becomes as narrow as 100 km
in Cte dIvoire, 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
Cte dIvoire
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 (%)
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, Ebri and Aby in Cte
dIvoire, and Keta in Ghana). The main water bodies on
the continent are formed by lakes Buyo and Kossou in
Cte dIvoire 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 Konkour in Guinea,
Scarcies, Moa and Mano in Sierra Leone, Lofa, St. Paul,
St. John and Cestos in Liberia, Cavally, Sassandra, Como
in Cte dIvoire, 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 Cte dIvoire, and Tonkui (1190 m)
in Cte dIvoire. 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
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
Soil class
cmk < 4 and ph < 5.5
cmk 4-8 and ph 5.5
cmk > 8 and ph 5.5
At Cape Palmas, at the Liberian-Ivorian border, the

coast deflects towards the northeast. The western part of
Cte dIvoire 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 24-28C. 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
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.
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 Cte dIvoire
consists mainly of granite and migmatite, with intrusions
of metamorphic schists near Ta National Parc (Avenard
1971). The eastern part of Cte dIvoire 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 FAOs 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 (Cte dIvoire 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 Cte dIvoire 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 Cte dIvoire
having a mixture of these (Figure 1.3E).
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Vegetation
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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Page 11
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 Cte dIvoire. 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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Page 12
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 Cte dIvoire,
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 Kouam et al. provide a detailed analysis
of the floristic diversity in Cte dIvoire. In Cte dIvoire
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 VBaoul. It has been hypothesised that this savanna
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Page 13
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 Guin-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, Kouam et al. discuss the role of the V-Baoul
and other environmental factors in determining species
composition of Ivorian forests.
The rainforests of West Africa have been earmarked
as one of the worlds 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

Cte dIvoire. 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 (Cte dIvoire) 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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Page 14
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.
10:24 AM
Page 15
Forest cover changes in

Cte dIvoire and Upper Guinea
C. Chatelain, H. Dao, L. Gautier and R. Spichiger
H A P T E R
11/11/03
22466 layout Chapter 2 p15-32
Introduction
Of the forest that covered West Africa at the

beginning of the 20th century, the large majority has
disappeared and the Ta National Park forest in Cte
dIvoire 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
forests 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 Cte dIvoire, a
country with a long history of forest exploitation (Meniaud
1922, Aubrville 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 Cte dIvoire has been
assessed at 7.6% per year (850 km2/yr). In 1990 only
11,230 km2 of dense forests were remaining in Cte
dIvoire, 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 forts classes
of Cte dIvoire 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
(Pivinen 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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Page 16
Chapter 2. Forest cover changes in Cte dIvoire and Upper Guinea
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 Cte d'Ivoire, including Ta 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 Cte dIvoire 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 Cte dIvoire in 1993 (Dao 1999) based
on NOAA-AVHRR images. The Ta forest area represents at least
40% of the total forest area of Cte d'Ivoire. Also the forest area south
of Abengourou (the classified forests of Yaya-Bossmati-Mabi) is very
important. A full-colour version of this figure can be found in
Appendix 1.
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Page 17
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 Cte dIvoire, 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 Cte
dIvoire. 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 Cte dIvoire, 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-Baoul (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 Cte dIvoire Geographical Institute (Dao
1999) which are based on aerial photographs taken in
1958 and which are more precise than Guillaumet and
Adjanohouns 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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10:25 AM
Page 18
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
Lovigui
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
Hir
Go
Abi
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.
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
11/11/03
10:25 AM
Page 19
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, Chevaliers 1912 map
(Chevalier 1920) shows an area of 29,205,000 ha in Cte
dIvoire, Liberia, Ghana and Sierra Leone (Table 2.3). The
same analysis of Shantzs 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
Cte 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
19
11/11/03
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Page 20
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 Cte dIvoire, a little under half of
Ghana (5,830,000 ha), and a little bit of Togo, today there
remains only the Ta National Park in Cte dIvoire, 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 Cte dIvoire 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 Pivinen 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 Cte
dIvoire, 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 Cte
dIvoire, which is considerable (Table 2.4). The forts
classes cover 2/3 of the protected areas, while national
parks represent 1/3. The forts classes 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, Como (in the northeast of the country,
1,167,167 ha) and Ta (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
forts classes (71%) have an area equal to or less than
National scale: Cte dIvoire
The oldest map showing forest cover of Cte dIvoire

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 Chevaliers 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 Cte dIvoire, 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 Cte
dIvoire 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 Ta
region with the Ta National Park, the Goin-Db and
Cavally forts classes; the Agboville-Abengourou region
with the forests of Mabi-Yaya and Yapo, and the Guitry
region with the G-Bodinou fort classe. At the same
time some regions have no forest at all. This is the case
most notably in the Daloa-Gagnoa-Soubr triangle in the
Central-West region, which was the first zone of
agricultural colonisation that moved towards the countrys
southwest during the 1960s.
Compared with Monniers (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 Cte dIvoire 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
11/11/03
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Page 21
variable area of protected forest cover. The West region of

Cte dIvoire (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 Ta forest. In the same
way, the Northeast region, which includes the Como
national park has 24.4%. At the scale of Cte dIvoire, the
regions most poorly covered with protected forests are
found in the borders of the North (around Tingrla, 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 forts classes (less than
10,000 ha), than the forest zone, that is, 66% compared
to 43%.
The actual forest cover
The different regions in Cte dIvoire 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 (%)
Ta (P.N.)
Hte Dodo
Cavally-Goin-Db
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.)
Nigr
Scio
Dukou
Mt-Pko (P.N.)
Okromodou-Diogoro-Bogbo
Go-Bodinou
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-Sangouin
Bki-Bossmati
Asagny (P.N.)
Mt Glo
Krozial
Dogodou
N'Guchi
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
Hir
G
Yapo
Abi
Sikensi
Lovigui
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
21
11/11/03
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Page 22
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 Ta, GoinDb-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 Cte 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-Soubr 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 Adzop (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 regions 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 forts classes and for the national
parks. The savanna zone shows an increase in both number
and area of forts classes. 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 Baoul peoples 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 forts classes of the forest zone, remnant
forests represent on average 42% of their area. The area of
these forts classes 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 fort classe, the Mabi-SonganTamin-Yaya grouping, and Beki-Bossmati.
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 forts classes were still little touched by
agriculture. The map of 1958 showed evidence of deforestation mostly in the east of the region (Adzop, 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
11/11/03
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Page 23
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 Tiassal-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 G-Bodinou 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 G-Bodinou 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 Tn 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
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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 forts classes. 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, G-Bodinou 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 (Hir region). Some forts classes
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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The Hir 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 forts classes 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 Hir (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 fort classe was completely transformed in oil palm plantations.
The G-Bodinou 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 G-Bodinou fort classe. The landscape
to the north of this forest is made up of numerous groves. Para-rubber tree and palm
plantations have increased heavily. This fort classe, along with those of Yapo and Niegr,
are the three last large areas of forest in the South zone of the country.
The Lovigu 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 forts classes. In 1969,
deforestation already reached the borders of the forts classes. In 1990, over 40% of the
forts classes of Maf and Lovigu were already cleared, and in the decade to 2000, most
clearing was done in the Maf fort classe. This block, along with that of Abi, shows the
highest forest cover in 2000, mostly situated in the rural domain.
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 fort classe 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 Abi block north of the Yapo fort classe. 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 forts classes (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.
25
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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 forts classes in Cte
dIvoire 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, LHote & Mah 1995).
In Cte dIvoire, 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.
Cte dIvoire: 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 forts classes. 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 forts classes, it has not
been simple to follow the development of these areas. In
fact forts classes 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
forts classes) 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 forts classes still exist today, often as groves
that are difficult to access or are unexploitable. In and
around forts classes 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 forts classes area has been cleared. Numerous
factors determine conservation: the existence of real
management and protection projects (as for example in
the Ta 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 Cte dIvoire, 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 Cte 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 G-Bodinou fort classe.
This supports our believe that the role of forts classes
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 Ta
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 G-Bodinou 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 forts
classes, 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 Cte dIvoire.
Figure 2.10 Changes in area under cultivation in Cte dIvoire. The

principal commercial cultures of the forest zone are given.
2.
Causes of deforestation
The causes of deforestation in Cte dIvoire 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 Cte dIvoire.
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-Baoul in the middle of Cte dIvoire,
(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
28
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 Cte dIvoire,
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 Cte dIvoire, 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 Cte dIvoire, most notably the savanna
regions (such as the Baoul), 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, Cte dIvoire had a low population density,
especially in the southern forests. The traditional agrarian
system of Cte dIvoire 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 Cte dIvoire 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), Cte dIvoire 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 Cte dIvoire
itself.
The role of the State: Besides the agrarian option
chosen by Cte dIvoire 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), Cte dIvoire 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 forts classes 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
(Ta, 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
Cte dIvoire, 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
29
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30
Figure 2.11 Production of wood logs in Cte dIvoire: comparison of

produced and exported log volumes.
Figure 2.12 Development of human population and of deforestation in

Cte dIvoire.
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 Cte
dIvoire 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 Cte dIvoire 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 Tiassal, Rubino, and
MBatto), 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: forts classes. It is urgent that
the government of Cte dIvoire develops a real policy of
conservation regarding the forts classes in the country,
even though only 23% of the remaining forest in the
country is within these forts classes. This study of the
development of deforestation shows that conserving forests
by the creation of forts classes 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 forts classes. 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 Unions 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 Ta region (Bonnhin, 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 Cte dIvoire will be the
principal actors in putting a conservation program into
place, as these countries have most of the remaining
forests.
31
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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 Cte dIvoire, 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 Ta 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 Cte
dIvoire 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, Cte dIvoire
possesses an important reservoir of protected areas. There
are numerous forts classes 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 forts classes they have played a role in the
conservation of the natural setting, because the most
cleared regions, such as the central regions of GagnoaSoubr or Daoukro, are characterised by the absence of
these protected areas.
32
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 Cte dIvoire 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
L. Gautier and R. Spichiger
H A P T E R
11/11/03
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-Baoul, central Cte dIvoire.
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. 8N,
with notable exceptions in central Cte dIvoire, where
savanna penetrates as far south as 6N in a region called
V-Baoul, 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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Chapter 3. The forest-savanna transition
B
Figure 3.2 Forest-savanna mosaic in Lamto, Cte dIvoire.
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 prforestier(Monnier 1981) or
the Guineo-Congolian/Sudanian transition zone (White
1983). This terminology underlies a very important
34
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 (Aubrville
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 Cte dIvoire 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 Cte dIvoire (e.g. in Dabou,
Grand-Bassam or Grand-Lahou) and the persistence of
rainforest outliers in the north of V-Baoul (e.g. in
Bamoro), indicate that the northern limit of the
continuous Guinean rainforest has been fluctuating during
the late Quaternary. Aubrville (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 (Aubrville 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. (Frdoux 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
B
Figure 3.3 The forest-savanna mosaic as seen from the ground in
Lamto, Cte dIvoire. 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 VBaoul down to the sea could have led to allopatric
speciation resulting in sister species on both sides of the
separation (see Kouam 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).
35
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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, Bourlire 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, Aubrville (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
Cte dIvoire, at the eastern side of the V-Baoul 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, Aubrville 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,
mans 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 Bouak in Cte dIvoire, 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-Baoul) 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 Adiopodoum, south Cte dIvoire, 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. 9N) 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 Aubrville
(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
C
Figure 3.4 The Kokondekro experiment (central Cte dIvoire) after 55
years. A) The early burning plot, B) the late burning plot, C) the
protected plot.
37
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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
(Aubrville 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 Cte dIvoire,
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).
Aubrville (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 Cte dIvoire (V-Baoul). 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-Baoul,
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-Baoul 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
11/11/03
10:26 AM
Page 39
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 (Aubrville 1966). This has been demonstrated by
comparing two sets of aerial photographs separated by
25 years in Lamto reserve, Cte dIvoire. 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-Baoul, 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 Cte dIvoire, 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.
39
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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
40
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.
10:27 AM
Page 41
H A P T E R
The forests of Upper Guinea:

gradients in large species composition
F. Bongers, L. Poorter and W.D. Hawthorne
11/11/03
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, Aubrville 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 Ta region, van Rompaey 1993 for southeast Cte
dIvoire - 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, Kouam 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 Ta National Park, from Mt Nienokou,

Cte dIvoire.
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.
41
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Chapter 4. The forests of Upper Guinea: gradients in large species composition
Methods
Study area, plot- and data selection
The area covered in this analysis is confined to the
forest zone of Sierra Leone, Liberia, Cte dIvoire 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
Cte dIvoire we drew largely on data collected by
SODEFOR (Clment & Guinaudeau 1973, SODEFOR
1978, 1979), and for some sites on species lists (Kouam 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
Cte dIvoire 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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10:27 AM
Page 43
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 Cte
dIvoire 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
Cte dIvoire 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 Cte dIvoire. 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 Wards 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
43
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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) Species. The abbreviations

refer to the species in Table 4.1.
(B) Sites. The symbols refer to
the country.
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 Pearsons 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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Page 45
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.
(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.
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 Cte dIvoire 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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10:27 AM
Page 46
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 Cte dIvoire 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 Pearsons r= 0.95, p<0.001; second
axis Pearsons 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 Cte
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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Page 47
dIvoire (WE3), in a band east of WE3, mainly in the

south west of Cte dIvoire (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 Cte dIvoire 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 Cte dIvoire, 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 Cte
dIvoire, 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 Cte dIvoire 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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Page 49
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.
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, Cte
dIvoire, 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
49
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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 Cte dIvoire. The map for Ghana is
based on Hall & Swaine (1976) and they extended that
system into Cte dIvoire. Wet evergreen forest was found
in the southeast and in the southwest of Cte dIvoire,
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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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 Cte
dIvoire, with between 1400 and 1500 mm rainfall per
year.
In Cte dIvoire, 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 Kouam 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
Cte dIvoire 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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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.
10:27 AM
Page 53
Floristic diversity of
closed forests in Cte dIvoire
F. N. Kouam, K.E. Kouadio, K. Kouassi and L. Poorter
H A P T E R
11/11/03
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 Cte dIvoire, the savanna
penetrates the forest zone in a region called V-Baoul
(Guillaumet & Adjanohoun 1971, Gautier & Spichiger
chapter 3). On each side of V-Baoul, 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, Ta), 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 Cte dIvoire (Aubrville
1949, Guillaumet 1967, Anonymous 1991, Beentje et al.
1994). Guillaumet and Adjanohoun (1971) have
distinguished a floristic species group in southwest Cte
dIvoire which they coined Sassandrian. This group
includes species that occur between the Cavally river and
the Sassandra river, from Tabou to Ta. Another species
group, called Ghanaian, occurs in southwest Ghana (Hall
& Swaine 1981), and appears to penetrate into the east of
Cte dIvoire.
In this chapter, we investigate the environmental
factors responsible for the floristic composition in the
lowland closed forests of Cte dIvoire. The forest blocks of
southeast and southwest Cte dIvoire are currently
separated by the relatively dry region of the V-Baoul.
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 Tr showing the beautiful

cauliforous tree Omphalocarpum elatum (Sapotaceae) in Banco
National Park, Cte dIvoire.
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 Cte dIvoire. 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
Bossmati
Bm
SD
22,200
6 20 - 6 35N
3 20 - 3 35W
1400 - 1500
5 51 - 6 05N
3 22 - 3 41W
1650 - 1700
5 46 - 6 12N
3 12 - 3 26W
1600 - 1650
5 40 - 5 58N
3 11 - 3 25W
1650 - 1700
5 35 - 5 54N
3 23 - 3 46W
1700 - 1800
5 00 - 5 07N
5 50 - 5 57W
1550 - 1600
4 44 - 4 58N
6 14 - 6 35W
1650 - 1750
5 06 - 5 11N
5 29 - 5 34W
1550 - 1600
5 21 - 5 25N
4 01 - 4 05W
2000
6 22 - 7 24N
6 59 - 7 10W
1460 - 1680
6 53 - 7 14N
5 46 - 6 10W
1400
4 41 - 5 19N
7 01 - 7 25W
1900 - 2300
5 09 - 6 09N
6 48 - 7 26W
1800 - 2200
Southeast
Mabi
Songan
Tamin
Yaya
South coast
Dassioko
Monogaga
Port Gauthier
Banco
Centre-West
Southwest
Mo
PG
Haut
Sassandra
HS
Marahou
Ma
Haute Dodo
Ta
1.
2.
3.
4.
Mb
Guillaumet & Adjanohoun (1969)

Eldin (1971)
Perraud & Souchre (1970), De Rham (1971)
Eldin & Daudet (1968)
HD
Ta
SD
SD
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
Ak 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
Ak Assi (1997)
Sand
300
859
Kouam (1998b)
Sand
450
705
Kouam (1998b)
Sand
200
773
De Koning (1983)
Granite
400
843
Kouam (1998a)
Schist
350 - 400
475
Jongkind et al. (1999)
Granite
150
906
Kouadio (2000)
Kouassi (2000)
Granite
200 - 250
849
Ak Assi & Pfeffer (1975)
Page 54
Forest type1
10:27 AM
Abbr.
11/11/03
Forest
Chapter 5. Floristic diversity of closed forests in Cte dIvoire
Geographical Zone
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.
11/21/03
2:02 PM
Page 55
Figure 5.2 Map of Cte dIvoire 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-Baoul.
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 Srensons 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
Srensons 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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10:27 AM
Page 56
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
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
Landolphia hirsuta
Landolphia owariensis
Microdesmis keayana
Myrianthus arboreus
Myrianthus libericus
Napoleonaea vogelii
Nephrolepis biserrata
Neuropeltis acuminata
Ochthocosmus africanus
Palisota hirsuta
Parinari excelsa
Psychotria peduncularis
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 Marahou (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 (Ta/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/Marahou) to
53% (Bossmati/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 (Bossmati, Haut
Sassandra, Marahou), 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, Ta), 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
11/11/03
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centres of greatest floristic diversity in the lowlands in

Cte dIvoire. 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 Pearsons 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 Srensens 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
Bossmati
Dassioko
Haut
Haute
Sassandra Dodo
Port
Gauthier
Mabi
Marahoue
Monogaga
Songan
Ta
Bossmati
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)
Ta
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
11/11/03
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Figure 5.3 Relationship between the number of species in common and

the number of forests compared. Spearmans 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 Cte dIvoire. The high
contribution can be explained by the fact that all principal
lowland forest types in Cte dIvoire 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 Cte
dIvoire 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
58
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 (Ak Assi 1984).

The Fabaceae, represented in the Guinean savannas by 103
species (Kouam 1993, Bnninger 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.
11/11/03
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Page 59
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
Bossmati 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 & Souchre 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 Cte dIvoire 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
Dougoun, Amadou Fofana and Patrice Maba for their
assistance in the data collection. We thank Profs Laurent
Ak Assi for identifying the taxa, and Dossahoua Traor
and Frans Bongers for their comments, which have
allowed us to improve this chapter.
59
60
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Biodiversity hotspots in
West Africa; patterns and causes
J.J. Wieringa and L. Poorter
H A P T E R
11/11/03
Introduction
The rainforests of West Africa have been earmarked as

one of the worlds 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 (Aubrville 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 Cte
dIvoire, 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, OBrien 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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Chapter 6. Biodiversity hotspots; patterns and causes
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 Cte
dIvoire (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
Cte dIvoire 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),
62
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 Cte dIvoire
present in the Geneva herbarium database (Conservatoire
et Jardin Botaniques de la Ville de Genve, Switzerland)
(only as far as the data was owned by Geneva), the
collections from Cte dIvoire and Guinea present in the
Paris herbarium database (Musum National dHistoire
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 Cte
dIvoire and Guinea. Since we have abundant collections
from Cte dIvoire 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, Cte dIvoire. 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 Ta, 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, Ta
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
64
Figure 6.4 Fitted species collection curves for 50 x 50 km areas

around Tabou (continuous line), Ta (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 Spearmans 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
(Aubrville 1949, Guillaumet 1967). To a lesser extent it
has been based on the occurrence of species with restricted
distribution ranges.
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6. Biodiversity hotspots in West Africa; patterns and causes
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
weight
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
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
65
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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 Adiopodoum 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 Tk forest, and the cell containing the
Ecological Station at Ta 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 Ta National Park and the Scio
66
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 Cte
dIvoire and then fading away towards Sassandra. The
climax of this range lies in Liberia and southwest Cte
dIvoire. 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 Cte dIvoire. 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 Como
National Park in Cte dIvoire (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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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
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.
68
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 LiberiaCte dIvoire 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 Spearmans 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
rs
rs
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
Variable
***
***
***
***
C. Palmas
Rain
Rain2
Mt Nimba
Rain x WHC
Slope
- 0.62
0.06
- 0.11
0.44
- 5 x 10-7
rs
0.29
0.35
0.38
0.42
0.44
***
***
***
***
*
69
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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 Cte dIvoire 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 (Kouam 1998a). The Como 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 (OBrien 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, OBrien 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
70
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 (OBrien 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
72
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.
Page 73
M. Holmgren, L. Poorter and A. Siepel
What explains the distribution of

rare and endemic West African plants?
10:29 AM
H A P T E R
11/11/03
Introduction
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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Chapter 7. Distribution of rare and endemic West African plants
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, Ak 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
74
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
present in the Geneva herbarium database (Conservatoire
et Jardin Botaniques de la Ville de Genve, Switzerland),
the collections from Cte dIvoire and Guinea present in
the Paris herbarium database (Musum National dHistoire
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
11/11/03
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Page 75
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 collectors 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 collectors
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 (Pearsons 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
75
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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
11/11/03
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Page 77
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
Cte dIvoire 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).
77
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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 Cte dIvoire and
Ghana, and a second population at the border between
Cte dIvoire and Liberia (Cassipourea hiotou) or further to
the west (Monocyclanthus vignei, Pierreodendron kerstingii,
Schumanniophyton problematicum, Strephonema
pseudocola). Both populations, especially the one between
Cte dIvoire 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=Cte dIvoire, 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
78
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
CdI
Gh
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
submontane
unclear
coastal
inland
coastal
unclear
coastal
inland
montane
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
11/11/03
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Page 79
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.
(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), Cte
dIvoire (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).
79
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Page 80
(average r2 = 0.27, range 0.050.63). 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.
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 (Pearsons r,
p < 0.001 in all cases), and also with the percentage of
records found in open places (Pearsons 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.
11/11/03
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Page 81
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 Cte dIvoire 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, Kouam 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
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
81
11/11/03
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Page 82
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
Sabicea bracteolata
Tarenna vignei var. vignei
Byttneria dahomensis
Byttneria ivorensis
Ancistrocladus pachyrrachis
Uvaria dinklagei
Suregada ivorensis
Leucomphalos libericus
Millettia leonensis
Ixora liberiensis
Keetia obovata
Sericanthe adamii
Byttneria guineensis
Premna grandifolia
82
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
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
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 Cte dIvoire and Ghana, at the border between
Cte dIvoire 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-Baoul, a
savanna intrusion in Cte dIvoire (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 Cte dIvoire, for example,
are clear vicariant species (Hall & Swaine 1981). The
importance of Cape Palmas (southwest Cte dIvoire) 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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(Aubrville 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 Cte dIvoire, 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-Cte dIvoire and the borders of Cte
dIvoire-Ghana (Fig. 2.2). For the Liberian-Cte dIvoire
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 Cte
dIvoire. 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 Ta National
84
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 Cte dIvoire
connect the semi-deciduous forests of Haut Sassandra

with the moist evergreen forests of Ta-NZo and Nigr
using the Sassandra river as a corridor.
For the Cte dIvoire-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

Cte dIvoire 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 Pko mountain range between Guinea,
Liberia and Cte dIvoire, 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.
85
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Implications for conservation

and management
F. Bongers, L. Poorter, V. Belign, W.D. Hawthorne, F. N. Kouam,

M.P.E. Parren and D. Traor
H A P T E R
11/11/03
Need for effective conservation and

sustainable use
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
Cte dIvoire, 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, Cte dIvoire, 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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Chapter 8. Implications for conservation and management
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 Cte
dIvoire and in southeast Liberia. Smaller, but numerous
88
tracts occur in west Ghana. Ta 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). Ta
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 Cte dIvoire. The detailed local analyses of
Chatelain et al. (chapter 2) show that the rural areas of
Cte dIvoire 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 Cte 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-Baoul,
in Cte d'Ivoire to the west. Movement of elephants between
forest fragments, often not exceeding 10,000 ha, in the Cte
dIvoire-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 Cte dIvoire
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
Bossemati (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. 9N) 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 (Aubrville 1966). This has been
demonstrated by comparing two sets of aerial photographs
separated by 25 years in Lamto reserve, Cte dIvoire.
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 Cte dIvoire 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
90
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 (Aubrville 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-Baoul (see Kouam 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 Cte dIvoire 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 Cte dIvoire.
The climax of this belt is in Liberia. Additional high
diversity areas are found in areas around Abidjan in Cte
dIvoire and Ankasa in Ghana. A second belt is found in
the mountain areas of Mt Nimba/Mt Ziama in Guinea,
and around Man in Cte dIvoire, 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 Cte dIvoire 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 Cte dIvoire
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 Cte dIvoire and the other is the
area in southwest Ghana (see Figures 1.3 and 4.3). In the
Liberian-Cte dIvoire 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 Ta forests, and between Haut
Sassandra and Ta-NZo and Nigr. In the Cte dIvoireGhana 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 Pko mountain
range between Liberia, Guinea and Cte dIvoire, 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 Cte dIvoire 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 Cte dIvoire from H. Dao and C. Chatelain, amended by V. Belign.
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.
92
A. Forest condition
Ghana
Cte dIvoire
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
Cte dIvoire
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
Cte dIvoire
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 Cte dIvoire Kouam 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. Kouam et al. (chapter 5)
distinguish four forest types with local endemic species.
Consequently, forest management and protection in Cte
dIvoire 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: Ta
NP for western evergreen forest, Banco NP for coastal and
Marahou 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 Cte-dIvoire 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 Ghanas 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 Cte dIvoire 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 Cte dIvoire, 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 Cte dIvoire,
Liberia and Guinea.
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Figure 8.2 A large Ceiba pentandra tree is cut in a forest concession at

NZo (Cte dIvoire).
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 Cte dIvoire
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, Cte dIvoire
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 Cte dIvoire, for instance, only
11 areas are formally protected as national parks or
natural reserves, with in total 1,874,800 ha: Azagny,
Banco, Como, Haut-Bandama, Iles Ehotil, Marahou,
Mt Pko, Mt Sangb, the complex Ta & NZo and Mt
Nimba (Bakarr et al. 2001). Only Como and Azagny
National Parks were considered to have a reasonable level
of administrative and management support, enough to
94
provide real protection for the parks. All others are

considered to have shortage of effective protection. For Ta
National Park & NZo Reserve effective protection is
doubtful, despite numerous international agencies
maintaining research projects in the area. Ta 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 Ta (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 Cte dIvoire this was
done by Hy Dao, Chatelain and amended by Belign. 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 Ta 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 Cte dIvoire 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 Cte
dIvoire 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 Cte dIvoire,
however, do have an important role to play with respect to
biodiversity: many forest species are confined to one or a
few forests only (Kouam 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 Cte dIvoire 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
95
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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, Cte dIvoire.
directed to forest elephants. They focus on the border areas

between Ghana and Cte dIvoire (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
96
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, Ta-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 Ta-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, Ta-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 Cte dIvoire (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
Waldrons red colobus) disappeared completely from
previously occupied localities in Ghana and Cte dIvoire
(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 Cte dIvoire.
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 Abbiws (1990) book. For two forests
in Cte dIvoire, 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 Cte dIvoire;
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 Cte dIvoire 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 Cte dIvoire 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 Cte dIvoire and
border with Liberia, the other in southwest Ghana
and border with Cte dIvoire.
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.
98
Ghana has most of its forest types sampled in protected

forests, in Cte dIvoire 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 Cte dIvoire 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.
22466 layout page 099-107
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B
Species
Chapters 9-11
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Page 101
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
10:32 AM
H A P T E R
11/11/03
Introduction
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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Chapter 9. Ecological profiles of rare and endemic species
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
present in the Geneva herbarium database (G:
Conservatoire et Jardin Botaniques de la Ville de Genve,
Switzerland) the collections from Cte dIvoire and Guinea
present in the Paris herbarium database (P: Musum
National dHistoire 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-Juams 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.).
102
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
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 collectors 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 (Pearsons 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 collectors 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, clairire, clearcut, clearing,
cultivated, degraded, dnude, disturbed, edge, farm,
fourr, fringe, fringing, gap, jachre, lisir, 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
104
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 todays 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,
105
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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
106
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 Cte
dIvoire from the Genve 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 Sonk (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 Kouam. 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
108
% 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, Cte
dIvoire, 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
Open (n)
Altitude
River
Coast
Mean Annual Rainfall
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.g.
28
25 (8)
11
39
43
32
25
50
50
39
H
21
All
46333
37
39
37
29
24
10
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)
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, Cte d'Ivoire,
Ghana
spp
110
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 Cte 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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.a.
27
100 (14)
41
37
15
78
26
41
33
19
30
H
41
All
46333
37
39
37
29
24
10
69
25
36
13
39
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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)
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
Upper Guinea: Senegal, Guinea-Bissau, Guinea,
Sierra Leone, Liberia, Cte dIvoire, Ghana
in 27 30 cells, distribution range is 1904 km
Forest type: wet forest, dry forest, secondary
forest, thickets
spp
Open (n)
Data sources
FWTA, Hawthorne & Jongkind (2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.p.
52
54 (24)
13
38
23
17
30
25
26
48
52
35
21
H
17
All
46333
37
39
37
29
24
10
69
25
36
13
39
111
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Adhatoda robusta
C.B.Clarke
Acanthaceae
A
Description
Phenology
Guild: np
Life form: shrub or small tree
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
Other: the bark outside is green with globular
lenticels. The sapwood is yellow-white.
Data sources
Distribution
Continent: Nigeria, Cameroon, Equatorial Guinea,
Gabon, Democratic Republic of Congo
Upper Guinea: Cte dIvoire, Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.r.
50 (4)
11
67
33
67
33
67
22
22
All
46333
37
39
37
29
24
10
69
25
36
13
39
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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

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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.a.
0 (7)
13
75
13
88
38
63
13
38
All
46333
37
39
37
29
24
10
69
25
36
13
39
113
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Afzelia parviflora
(Vahl) Hepper
Leguminosae-Caes.
A
Description
Habitat
Guild: np
Life form: medium-sized tree
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
dIvoire
Forest type: wet evergreen forest, savanna, gallery
forest, coastal shrubland, secondary forest
Data sources
Hawthorne & Jongkind (2004)
spp
114
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
A.p.
27
50 (20)
41
15
26
15
53
93
70
H
15
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:32 AM
Page 115
Alafia parciflora
Stapf
Apocynaceae
Description
Phenology
Guild: pi
Life form: winding woody climber
Max. height: 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
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.p.
100 (2)
100
33
67
100
67
All
46333
37
39
37
29
24
10
69
25
36
13
39
115
11/11/03
10:32 AM
Page 116
Albertisia cordifolia
(Mangenot & Mige) F
Menispermaceae
A
Distribution
Data sources
Continent: Upper Guinea endemic (De Koning

1983)
Upper Guinea: Cte dIvoire
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

Timing: flowering period from April to June; fruiting
period from June to July and January to February
(De Koning 1983)
Guild: np
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.c.
14
9 (11)
100
93
21
79
21
79
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:33 AM
Page 117
Albertisia cuneata
(Keay) Forman
Menispermaceae
A
Description
Phenology
Guild: sb
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.
Data sources
Troupin (1962), Hall & Swaine (1981)
Distribution
Upper Guinea: Senegal, Cte dIvoire, Ghana. It
was originally described for Ghana, but has been
subsequently found in Senegal (Hall & Swaine 1981)
and Cte dIvoire (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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.c.
0 (4)
25
75
25
100
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
117
11/11/03
10:33 AM
Page 118
Albertisia ferruginea
(Diels) Forman
Menispermaceae
A
Description
Data sources
Guild: u
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),
Distribution
Upper Guinea: Sierra Leone, Liberia
Distribution type: continuous, local, present in 4
30 cells, distribution range is 274 km
Habitat
Data unavailable.
Phenology
spp
118
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.f.
100 (2)
43
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:34 AM
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)
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.
Guillaumet & Debray (1964), Hawthorne & Jongkind

(2004)
Distribution
Distribution type: present in 2 30 cells.
Guillaumet & Debray (1964) mention at least 8
collections for Cte dIvoire.
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
3
46333
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
100 (1)
100
33
66
33
67
33
37
39
37
29
24
10
69
25
36
13
39
119
11/11/03
10:34 AM
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)
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

Phenology
Timing: flowering period from June to July; fruiting
period throughout the year (De Koning 1983)
Distribution
Upper Guinea: Liberia, Cte dIvoire, Ghana
Forest type: In Ghana, it is found in a wide variety
spp
120
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.s.
38
35 (23)
71
13
32
68
37
55
29
47
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:34 AM
Page 121
Alsodeiopsis chippii
Hutch.
Icacinaceae
Description
Phenology
Guild: sb
Max. height: 1.8 m (herbarium)
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

Upper Guinea: Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
17
83
33
67
33
67
24
10
69
25
36
13
39
A.c.
0 (6)
33
All
46333
37
39
37
29
Soil CMK
Soil WHC
121
11/11/03
10:34 AM
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
Other: it has a large and flat tuber.
Distribution
Continent: Burkina Faso, Benin, Nigeria
(Hetterscheid)
Upper Guinea: Sierra Leone (Hetterscheid), Cte
dIvoire, Ghana, Togo (herbarium)
Forest type: data unavailable
Habitat
Data unavailable.
Phenology
Description
Guild: u
Life form: large tuberous perennial herb
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
Open (n)
Data sources
Hetterscheid
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.b.
- (0)
100
75
25
50
25
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:34 AM
Page 123
Ancistrocladus pachyrrachis
Airy Shaw
Ancistrocladaceae
A
Description
Data sources
Guild: u
Life form: woody climber
Max. height: 3 m long (FWTA)
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.
Distribution
Upper Guinea: Liberia
in 1 30 cell
Habitat
Data unavailable.
Phenology
spp
Open (n)
A.p.
- (0)
All
46333
37
Altitude
River
Coast
>500m
<2km
<5km
VW
100
100
100
100
39
37
29
24
10
69
25
36
13
39
0
4
Soil CMK
Soil WHC
123
11/11/03
10:34 AM
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.

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
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
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
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, Aubrville (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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.l.
10
17
(6) 10
60
30
40
30
40
60
20
50
H
10
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:34 AM
Page 125
Anisophyllea meniaudii
Aubrv. & Pellegr.
Anisophylleaceae
Description
Phenology
Guild: np
Max. height: 21 m (FWTA)
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, Aubrville (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, Cte dIvoire,
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.m
19
14 (14)
37
58
21
21
68
32
37
63
All
46333
37
39
37
29
24
10
69
25
36
13
39
125
11/11/03
10:34 AM
Page 126
Anisotes guineensis
Lindau
Acanthaceae
A
Description
Data sources
Guild: u
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
FWTA
Distribution
Upper Guinea: Guinea
Forest type: wet evergreen forest, gallery forest
Habitat
It is found by streams (FWTA).
Phenology
spp
126
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.g.
- (0)
20
20
40
40
20
20
80
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:34 AM
Page 127
Anthocleista nobilis
G.Don
Gentianaceae
A
Description
Guild: pi
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
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
Upper Guinea: Senegal, Guinea, Sierra Leone,
Liberia, Cte dIvoire, Ghana
Forest type: rainforest, secondary forest, swamp
forest, gallery forest
Data sources
Regeneration
FWTA, Leeuwenberg (1961), De la Mensbruge

(1966), Jenk (1971), Hawthorne (1981, 1995a),
De Koning (1983), Kasparek (2000)
It has a phanerocotylar epigeal foliaceous seedling

type (cf. de la Mensbruge 1966).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.n.
52
70 (47)
15
46
12
12
33
36
18
15
60
25
33
17
H
25
All
46333
37
39
37
29
24
10
69
25
36
13
39
127
11/11/03
10:34 AM
Page 128
Anthonotha explicans
(Baill.) J.Lonard
Leguminosae-Caes.
A
Description
Phenology
Data sources
Guild: u
Life form: large tree
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
Liberia, Cte dIvoire
in 8 30 cells
Forest type: moist evergreen forest, savannawoodland
Habitat
It grows in dry places (herbarium).
spp
128
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.e.
60 (5)
11
67
22
11
11
11
66
56
44
33
H
44
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:35 AM
Page 129
Anthonotha vignei
Hoyle
Leguminosae-Caes.
A
Description
Phenology
Guild: np
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
Ghana
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
A.v.
56
50 (32)
55
18
34
39
26
82
11
39
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
129
11/11/03
10:35 AM
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
Ghana (doubtful record) (herbarium)
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
Leaf: alternate, simple, elliptic, macrophyll (4-16 x
12-39 cm), entire, glabrous; petiole up to 20 cm
long, pulvinate at the apex, glabrous
axis (spadix, up to 10 cm long)
spp
130
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.a.
0 (1)
20
20
20
40
60
40
60
40
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:35 AM
Page 131
Aphanocalyx microphyllus
(Harms) Wieringa ssp.
compactus
(Hutch. ex Lane-Poole) Wieringa
Leguminosae-Caes.
A
Description
Habitat
Guild: sb
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, Cte dIvoire
(Wieringa 1999)
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).
Phenology
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.m
21
0 (6)
33
48
48
71
29
71
10
All
46333
37
39
37
29
24
10
69
25
36
13
39
131
11/11/03
10:35 AM
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
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
Upper Guinea: Sierra Leone, Liberia (Wieringa
1999)
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
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.p.
20
0 (2)
55
20
100
85
15
70
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:35 AM
Page 133
Apodiscus chevalieri
Hutch.
Euphorbiaceae
A
Description
Data sources
Guild: u
Life form: small tree
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
Distribution
Upper Guinea: Guinea, Liberia, Ghana
Habitat
It is commonly observed on riverbanks on sandy
soils (herbarium).
Phenology
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.c.
0 (3)
86
29
71
86
14
71
14
All
46333
37
39
37
29
24
10
69
25
36
13
39
133
11/11/03
10:35 AM
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)
Leaf: opposite, simple, elliptic, microphyll (2.5-4 x
4-10 cm), entire, coriaceous, glabrous; interpetiolar
stipules
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)
dIvoire, Ghana (herbarium, Robbrecht 1981a)
Distribution type: continuous, widespread, in
Upper Guinea present in 22 30 cells. Rare in Ghana
(Hall & Swaine 1981).
forest, dry semi-deciduous forest, secondary forest,
thickets
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.a.
48
54 (28)
65
16
44
33
10
79
10
46
H
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:35 AM
Page 135
Argocoffeopsis lemblinii
(A.Chev.) Robbr.
Rubiaceae
A
Description
Phenology
Guild: u
Life form: shrub
Max. height: 1 m (Hawthorne & Jongkind 2004)
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.
Timing: only one record is known, and this
specimen flowered in January (FWTA).
Data sources
FWTA, Robbrecht (1981a), Hawthorne & Jongkind
(2004)
Distribution
Upper Guinea: Cte dIvoire (Robbrecht 1981a)
in 1 30 cell. Very rare; only known from the type
specimen Chevalier 17180 from Valle de lAgniby,
Voqui.
Forest type: moist evergreen forest
Habitat
The single record of this species was found in the
forest.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
A.l.
- (0)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
135
11/11/03
10:35 AM
Page 136
Asystasia scandens
(Lindl.) Hook.
Acanthaceae
A
Phenology
Uses
Sometimes it is cultivated as an ornamental plant
(FWTA).
Data sources
FWTA
Distribution
dIvoire
Description
Guild: u
Life history: woody climber
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
spp
136
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
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
A.s.
23
75 (4)
35
17
34
60
61
39
43
H
13
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:35 AM
Page 137
Baissea zygodioides
(K.Schum.) Stapf
Apocynaceae
Description
Phenology
Guild: np
Life form: large woody winding climber
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)
Data sources
FWTA, Hall & Swaine (1981), De Koning (1983), Van
Dilst (1995)
Distribution
dIvoire, Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.z.
59
60 (15)
32
40
23
26
10
53
44
36
H
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
137
11/11/03
10:35 AM
Baphia spathacea
Page 138
Hook.f. ssp.
spathacea
Leguminosae-Pap.
Distribution
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.
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
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
Open (n)
Data sources
FWTA, Soladoye (1984), Lock (1989), Hawthorne &
Jongkind (2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.s.
28
75 (16)
57
46
85
100
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:37 AM
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.
Distribution
dIvoire, Ghana
Forest type: (sub) montane rainforest
Phenology
Deciduousness: most leaves are shed in the dry
season
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
B.c.
25
40 (5)
32
36
24
48
28
72
28
48
32
All
46333
37
39
37
29
24
10
69
25
36
13
39
139
11/11/03
10:37 AM
Page 140
Begonia fusicarpa
Irmsch.
Begoniaceae
Description
Guild: u
Life form: perennial epiphytic herb
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)
Distribution
in 1 30 cell. It is an extremely rare species. Only
known from the type specimen J.T. Baldwin 11417
from Sino County, Kulo.
Habitat
Confined to lowlands in Liberia under 500 m altitude
(De Wilde 1983).
Phenology
Data unavailable.
Data sources
De Wilde (1983)
spp
140
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.f.
- (0)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:37 AM
Page 141
Begonia hirsutula
Hook.f.
Begoniaceae
Description
Guild: sb
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.
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.h.
- (0)
33
67
33
67
33
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
141
11/11/03
10:37 AM
Page 142
Begonia mildbraedii
Gilg
Begoniaceae
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)
Data sources
Sosef (1994)
Distribution
Continent: Cameroon, Gabon, Congo (Brazzaville),
Democratic Republic of Congo (Sosef 1994)
Upper Guinea: Cte dIvoire, 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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.m.
- (0)
60
100
20
40
40
20
20
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:37 AM
Page 143
Begonia prismatocarpa
Hook. ssp.
petraea
(A.Chev.) Sosef
Begoniaceae
Description
Phenology
Guild: sb
Max. height: 0.1 m
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
Other: this is one of three subspecies. The plant is
covered with minute glandular hairs.
Data sources
Distribution
Sosef (1994)
in only 1 30 cell. This is one of three subspecies,
and extremely rare; only known from the type
specimen Chevalier 19596 found between
Nkaougni 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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.p.
0 (1)
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
143
11/11/03
10:37 AM
Page 144
Begonia quadrialata
Warb. ssp.
nimbaensis
Sosef
Begoniaceae
Distribution
Upper Guinea: Guinea, Liberia, Cte dIvoire
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
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
Guild: sb
Max. height: 0.3 m
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
Other: the plant is covered with minute glandular
hairs.
spp
144
Open (n)
Data sources
Sosef (1994)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.q.
17
0 (3)
65
18
100
76
24
71
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:37 AM
Page 145
Beilschmiedia caudata
(Stapf ) A.Chev.
Lauraceae
Description
Data sources
Guild: u
Life form: tree
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
Other: the leaves, bark and stem have oil glands
and are sweetly scented.
FWTA, De Koning (1983), Mabberley (1987),

Distribution
Habitat
Data unavailable.
Phenology
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.c.
- (0)
33
17
51
83
17
83
17
All
46333
37
39
37
29
24
10
69
25
36
13
39
145
11/11/03
10:37 AM
Page 146
Beilschmiedia chevalieri
Robyns & R.Wilczek
Lauraceae
Description
Data sources
Guild: u
Life form: small to medium-sized tree
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
Other: the leaves, stem and bark have oil glands
and are sweetly scented.
FWTA, Mabberley (1987), Hawthorne & Jongkind

(2004)
Distribution
Upper Guinea: Cte dIvoire, Liberia
Forest type: wet evergreen forest, forest remnant,
secondary forest
Habitat
Data unavailable
Phenology
spp
146
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.c.
17 (6)
11
56
67
22
11
89
11
89
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:37 AM
Page 147
Berlinia occidentalis
Keay
Leguminosae-Caes.
Description
Guild: np
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).
Regeneration
Phenology
Dispersal: explosive
Timing: flowering period from March to May;
fruiting period from October to November (Taylor
1960)
Distribution
Ghana
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
Open ( n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
B.o.
41
38 (16)
44
20
30
25
38
10
90
56
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
147
11/11/03
10:37 AM
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
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.
Phenology
Dispersal: explosive pods (Hall & Swaine 1981)
Timing: not clearly seasonal (Hall & Swaine 1981)
Distribution
Data sources

Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.t.
37
73 (15)
49
14
44
25
19
62
32
30
H
43
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:39 AM
Page 149
Bertiera spicata
(P.Gaertn.) Wernham
Rubiaceae
Description
Guild: np
Life form: shrub
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
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).
Phenology
Distribution
Sierra Leone, Liberia, Cte dIvoire
Forest type: coastal forest, coastal shrubland,
mangrove forest, swamp forest, gallery forest,
secondary forest
Data sources
Habitat
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
B.s.
50
52 (21)
36
26
22
56
82
16
50
14
14
All
46333
37
39
37
29
24
10
69
25
36
13
39
149
11/11/03
10:39 AM
Bonamia vignei
Page 150
Hoyle
Convolvulaceae
Description
Phenology
Guild: u
Life form: large winding woody climber
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.
Data sources
FWTA, Hoyle (1934), Hall & Swaine (1981), Breteler
(1992)
Distribution
Upper Guinea: Ghana
Forest type: moist semi-deciduous forest, dry
forest, thickets
Habitat
Found in forest.
spp
150
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.v.
0 (7)
25
76
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:39 AM
Page 151
Brachystegia leonensis
Burtt Davy & Hutch.
Leguminosae-Caes.
Description

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).
Guild: sb
leaflets, elliptic, microphyll (0.5-4.5 x 1.5-12 cm),
entire, fairly coriaceous, young leaves densely
pilose, later glabrescent
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.
Habitat
It occurs on ridges and in valleys near water
(Hawthorne 1995a).
Regeneration
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,
Cte dIvoire
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.l.
20
11 (9)
60
50
50
75
20
35
35
All
46333
37
39
37
29
24
10
69
25
36
13
39
151
11/11/03
10:39 AM
Page 152
Buforrestia obovata
Brenan
Commelinaceae
Description
Phenology
Guild: sb
Life form: stoloniferous herb
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)
Distribution
Data sources
Continent: Upper Guinea endemic (herbarium)

Ghana (herbarium)
Forest type: locally abundant in lowland rainforest.
FWTA
Habitat
It is found on moist to damp forest floors
(herbarium).
spp
152
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
B.o.
0 (4)
25
38
13
38
50
13
75
25
63
H
25
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:39 AM
Page 153
Bussea occidentalis
Hutch. ex Chipp.
Leguminosae-Caes.
Description
Habitat
Guild: np
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
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).
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
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)

Ghana
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),
spp
B.o.
All
60
46333
Open (n)
22 (41)
37
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
42
20
45
23
68
32
43
42
39
37
29
24
10
69
25
36
13
39
153
11/11/03
10:39 AM
Page 154
Byrsanthus brownii
Guill.
Flacourtiaceae
Description
Phenology
Guild: u
Life form: small tree or shrub
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)
Other: the twigs are reddish with very conspicuous
small pale lenticels.
Data sources
FWTA, Berhaut (1975), Hawthorne & Jongkind
(2004)
Distribution
Continent: Upper Guinea endemic.
Upper Guinea: Senegal, Gambia, Guinea Bissau,
Guinea, Sierra Leone
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
Open (n)
B.b.
26
20 (5)
All
46333
37
Altitude
River
Coast
>500m
<2km
<5km
VW
58
19
27
12
54
62
38
19
27
27
39
37
29
24
10
69
25
36
13
39
Soil CMK
Soil WHC
H
11/11/03
10:39 AM
Byttneria dahomensis
Page 155
N.Hall,
B.guineensis, B.ivorensis Keay & Milne-Redh.

Sterculiaceae
Description
Habitat
Guild: u
Life form: prickly woody climber
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
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 (Hall
1962).
Distribution
Hall (1962), IUCN Red List (2000), Hawthorne

& Jongkind (2004)
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 Cte dIvoire. It has a continuous,
very local distribution, and is present in 1 30 cell. It
is a Red List species (extinct).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
Byttneria spp.
- (0)
75
50
50
75
25
50
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
155
11/11/03
10:39 AM
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.
Phenology
Timing: not distinctly seasonal (Hall & Swaine 1981)
Distribution
dIvoire, Ghana
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
Leaf: opposite, simple, elliptic to obovate,
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
Open (n)
C.s.
166
31 (107)
All
46333
37
Regeneration
It regenerates under closed canopy (Hall & Swaine
1981).
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
52
49
25
19
70
25
43
41
39
37
29
24
10
69
25
36
13
39
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)
11/11/03
10:39 AM
Page 157
Calpocalyx aubrevillei
Pellegr.
Leguminosae-Mim.
Desription
Regeneration
Guild: sb
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.

type (cf. Voorhoeve 1965). A gregarious tree, often
locally abundant (herbarium). Regeneration
needs light for healthy development
(Voorhoeve 1965).
Phenology
Dispersal: by explosive pods. Seeds are
eaten by chimpanzees (Kasparek 2000).
Timing: fruits are found the whole year round
(Voorhoeve 1965)
Distribution
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
C.a.
21
19 (16)
33
29
15
57
95
90
10
All
46333
37
39
37
29
24
10
69
25
36
13
39
157
11/11/03
10:39 AM
Page 158
Calpocalyx brevibracteatus
Harms
Leguminosae-Mim.
Distribution
Phenology
Continent: Sierra Leone to Nigeria (Hall & Swaine

1981), Nigeria, Cameroon (herbarium)
Ghana (herbarium)
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

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
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
(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
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
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.b.
62
33 (33)
34
47
21
23
66
29
44
52
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:41 AM
Page 159
Calvoa monticola
A.Chev. ex Hutch. & Dalziel
Melastomataceae
Description
Phenology
Guild: sb
Life form: small epiphytic or terrestrial herb
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
Distribution
Data sources
Continent: Cameroon, Gabon

Upper Guinea: Guinea, Liberia, Cte dIvoire,
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.m.
19
0 (6)
26
47
32
58
10
89
11
58
H
32
All
46333
37
39
37
29
24
10
69
25
36
13
39
159
11/11/03
10:41 AM
Page 160
Calycobolus insignis
(Rendle) Heine
Convolvulaceae
Distribution
Distribution type: continuous, local, present in
Forest type: mature forest, old secondary forest
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
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
Other: the stem has brown hairs.
spp
C.i.
All
160
Open (n)
Data sources
FWTA
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
33 (3)
67
17
84
100
83
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:41 AM
Page 161
Campylospermum amplectens
(Stapf ) Engl.
Ochnaceae
Description
Guild: sb
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
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
Other: it has aerial roots that sometimes penetrate
the leaf litter, and may help with nutrition on the
infertile evergreen forest soils (Hawthorne 1995a).
Regeneration
It regenerates in shade (Hall & Swaine 1981).
Distribution
Phenology

Ghana
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
Open (n)
Data sources
FWTA, Hall & Swaine (1981), Hawthorne (1995a)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.a.
16
10 (10)
88
31
63
100
69
31
All
46333
37
39
37
29
24
10
69
25
36
13
39
161
11/11/03
10:41 AM
Page 162
Campylospermum subcordatum
(Stapf ) Engl.
Ochnaceae
Distribution
Phenology

Forest type: montane forest, wet evergreen forest,
swamp forest, secondary forest.
Habitat
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
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
Flowers: medium-sized; calyx red; corolla bright
yellow
Fruit: fleshy, black on an enlarged red calyx;
1 seed
spp
162
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
C.s.
35
17 (23)
11
43
34
17
45
86
14
74
H
23
All
46333
37
39
37
29
24
10
69
25
36
13
39
Data sources
FWTA
11/11/03
10:41 AM
Page 163
Cassipourea afzelii
(Oliv.) Alston
Rhizophoraceae
Description
Phenology
Guild: sb
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
Distribution
Data sources
Continent: Nigeria
dIvoire, 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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
23
29 (7)
43
26
21
69
87
13
70
46333
37
39
37
29
24
10
69
25
36
13
39
163
11/11/03
10:41 AM
Page 164
Cassipourea hiotou
Aubrv. & Pellegr.
Rhizophoraceae
Description
Phenology
Guild: sb
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
Other: it has hairy twigs and petioles and a redbrown slash.
Data sources
Aubrville & Pellegrin (1958), Hall & Swaine (1981),
Hawthorne (1995a), IUCN Red List (2000)
Distribution
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)
Habitat
Found in the lower strata of rainforests.
spp
164
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.h.
100 (1)
67
33
66
100
67
33
all
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:41 AM
Page 165
Cathormion rhombifolium
(Benth.) Keay
Leguminosae-Mim.
Description
Phenology
Guild: u
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
Data sources
Distribution
FWTA, Brenan & Brummitt (1965), Voorhoeve


Upper Guinea: Senegal (FWTA), Guinea Bissau,
Guinea, Sierra Leone (herbarium)
Forest type: swamp forest, gallery forest
Habitat
Found in swamp forest and along riversides
(herbarium, Hawthorne & Jongkind 2004).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.r.
33 (3)
71
14
14
14
72
71
29
29
14
57
All
46333
37
39
37
29
24
10
69
25
36
13
39
165
11/11/03
10:41 AM
Page 166
Cavacoa baldwinii
(Keay & Cavaco) J.Lonard
Euphorbiaceae
Distribution
forest
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
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
spp
166
Open (n)
Data sources
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.b.
11
17 (6)
45
91
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:41 AM
Page 167
Chassalia corallifera
(A.Chev. ex De Wild.) Hepper
Rubiaceae
Description
Phenology
Guild: sb
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
Dispersal: probably by animals.

Distribution
Data sources
Continent: Upper Guinea endemic (Hall & Swaine

1981)
(herbarium)
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.c.
35
33 (21)
43
80
43
51
29
60
All
46333
37
39
37
29
24
10
69
25
36
13
39
167
11/11/03
10:41 AM
Page 168
Chazaliella cupulicalyx
Verdc.
Rubiaceae
Description
Phenology
Guild: u
Life form: scandent shrub
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
Other: it has a corky bark.
Data sources
Distribution
Verdcourt (1977), Hawthorne & Jongkind (2004)

Upper Guinea: Liberia (herbarium, Hawthorne &
Jongkind 2004)
Forest type: moist evergreen forest, secondary
forest
Habitat
Sometimes found near rivers (herbarium).
spp
168
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.c.
0 (2)
60
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:42 AM
Page 169
Chytranthus cauliflorus
(Hutch. & Dalziel) Wickens
Sapindaceae
Description
Guild: sb
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.
Distribution
Phenology
Continent: Gabon
forest, secondary forest
Habitat
In Ghana, its distribution is very highly correlated
with acidic, base-poor soils (Hall & Swaine 1981).
Regeneration
Data sources
FWTA, Hall & Ak Assi (1962), Aubrville & Leroy

(1973b), Hall & Swaine (1981), de Koning (1983)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.c.
12
11 (9)
33
67
25
33
58
83
All
46333
37
39
37
29
24
10
69
25
36
13
39
169
11/11/03
10:42 AM
Page 170
Jongkind
Verbenaceae
Habitat
Data unavailable
Phenology
Description
Guild: u
Life form: small creeping shrub
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
Other: most branches creep on the forest floor and
root
Data sources
Distribution
in 2 30 cells, distribution range is 10 km. It is only
known from southwest Cte dIvoire
spp
170
Open (n)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.s.
100 (1)
100
50
50
50
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:43 AM
Page 171
Cola attiensis
Aubrv. & Pellegr.
Sterculiaceae
Description
Guild: sb
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
Distribution
Phenology

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, Aubrville (1959), IUCN Red List (2000),
spp
Open (n)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.a.
14
0 (10)
29
57
43
79
14
43
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
171
11/11/03
10:43 AM
Cola boxiana
Page 172
Brenan & Keay
Sterculiaceae
Description
Phenology
Guild: sb
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
Distribution
Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
14 (7)
50
88
13
38
63
38
38
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:43 AM
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
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
Distribution
Ghana
Distribution type: continental disjunct, with the
largest population in Upper Guinea, present in 29
30 cells in Upper Guinea.
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
Open (n)
Data sources
FWTA, Hall & Swaine (1981), De Koning (1983),
Hawthorne (1994, 1995a), Swaine (1996)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.c.
71
38 (37)
52
21
31
32
14
11
68
21
34
13
H
52
All
46333
37
39
37
29
24
10
69
25
36
13
39
173
11/11/03
10:43 AM
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).
Habitat
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
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)

Distribution
Upper Guinea: Guinea (FWTA), Liberia, Cte
dIvoire, Ghana (herbarium)
spp
174
Open (n)
Data sources
Hawthorne (1995a), IUCN Red List (2000),
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.r.
24
0 (14)
33
42
67
25
54
46
25
58
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:43 AM
Page 175
Cola umbratilis
Brenan & Keay
Sterculiaceae
Description
Guild: sb
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
Distribution
Phenology

Upper Guinea: Ghana
in 3 30 cells, distribution range is 68 km, Red List
species (vulnerable)
Other: given its distribution, it is probably also
present in Cte dIvoire.
Habitat
It is found in forest understorey.
Regeneration
Data sources
FWTA, Hall & Swaine (1981), IUCN Red List (2000)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.u.
10
10 (10)
20
10
80
20
70
30
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
175
11/11/03
10:43 AM
Page 176
Combretum bipindense
Engl. & Diels
Combretaceae
Description
Phenology
Guild: np
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.
(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, Cte dIvoire, 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
Open (n)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.b.
10
0 (2)
40
10
60
30
10
10
50
40
30
30
30
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:43 AM
Page 177
Combretum blepharopetalum
Wickens
Combretaceae
Description
Phenology
Guild: u
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
Other: not much is known from this species, all
collections in the herbarium lack plant and
vegetation descriptions.
Dispersal: by wind
Data sources
Distribution
Wickens (1971), Mabberley (1987), Hawthorne &

Jongkind (2004)

Upper Guinea: Sierra Leone
Habitat
Data unavailable.
spp
C.b.
All
46333
Open (n)
- (0)
37
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
25
100
75
25
25
50
39
37
29
24
10
69
25
36
13
39
177
11/11/03
10:43 AM
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.
Distribution
Upper Guinea: Liberia, Cte dIvoire
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)
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
Open (n)
Data sources
FWTA, Mabberley (1987), Jongkind (in prep)
Altitude
River
Coast
>500m
<2km
5km
VW
Soil CMK
L
Soil WHC
L
C.c.
100 (3)
13
13
13
76
88
13
88
13
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:44 AM
Page 179
Combretum grandiflorum
G.Don
Combretaceae
Description
Phenology
Guild: pi
Max. height: over 20 m long (herbarium)
7.5-15 cm), entire, coriaceous, glabrous, shiny dark
green above, light green below, reddish veins,
prominent below
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
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
Data sources

Upper Guinea: Gambia, Senegal, Guinea Bissau,
Guinea, Sierra Leone, Liberia, Cte dIvoire, Ghana
Forest type: wet evergreen forest, swamp forest,
coastal shrubland, secondary forest
Habitat
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
C.g.
72
70 (30)
39
18
18
41
21
18
53
44
31
10
H
47
All
46333
37
39
37
29
24
10
69
25
36
13
39
179
11/11/03
10:44 AM
Page 180
Combretum tarquense
J.J.Clark
Combretaceae
Description
Guild: pi
Max. height: 3.5 m long (herbarium)
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
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 Cte

dIvoire and southwest Ghana.
rainforest, thickets
Habitat
It is often found in disturbed habitats; secondary
forest, forest edges, roadside thickets (herbarium).
Phenology
Dispersal: by wind
Distribution
Data sources

30 cells, distribution range is 159 km. It has a
spp
180
Open (n)
C.t.
11
60 (5)
All
46333
37

(2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
55
73
27
82
18
18
82
39
37
29
24
10
69
25
36
13
39
0
4
Soil CMK
Soil WHC
11/11/03
10:44 AM
Page 181
Combretum zenkeri
Engl. & Diels
Combretaceae
Description
Phenology
Guild: pi
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
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
Data sources
(2004)
Distribution
Continent: Benin, Nigeria, Cameroon
Upper Guinea: Guinea, Cte dIvoire, Ghana, Togo,
Benin
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.z.
47
75 (12)
30
81
19
38
62
26
47
All
46333
37
39
37
29
24
10
69
25
36
13
39
181
11/11/03
10:44 AM
Page 182
Commelina macrosperma
J.K.Morton
Commelinaceae
Description
Phenology
Guild: sb
Life form: perennial decumbent herb
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
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, Cte dIvoire, Ghana
(herbarium)
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
spp
182
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.m.
0 (3)
33
11
78
11
33
67
22
11
56
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:44 AM
Page 183
Commiphora dalzielii
Hutch.
Burseraceae
Description
Guild: pi
Leaf: alternate, compound, 3 leaflets, obovate to
rhomboid, microphyll (1.5-3 x 2.5-7 cm), entire,
herbaceous, glabrous
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.
Distribution
Upper Guinea: Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.d.
100 (9)
22
100
56
44
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
183
11/11/03
10:45 AM
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.
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
Upper Guinea: Guinea Bissau, Guinea, Sierra
Leone, Liberia, Cte dIvoire, Ghana
List species (vulnerable). It is a rare species.
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
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
(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
Open (n)
Uses
The bark is stripped from the tree, ground, and
used as a perfume (Voorhoeve 1965).
Regeneration
Data sources

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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.s.
23
0 (13)
43
39
34
17
26
52
22
43
17
H
26
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:45 AM
Page 185
Cordia vignei
Hutch. & Dalziel
Boraginaceae
Description
Phenology
Guild: pi
Leaf: alternate, simple, oblong to lanceolate,
mesophyll (5 x 20 cm), entire, lower surface with
long orange hairs, upper surface with white spots
Flower: medium-sized; calyx green; corolla white
with reflexed lobes; fragrant
Fruit: fleshy, ovoid (1.5 x 2 cm), yellow-green;
1-4 seeds
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),

Continent: Nigeria
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.v.
12
60 (5)
50
50
41
58
42
17
58
All
46333
37
39
37
29
24
10
69
25
36
13
39
185
11/11/03
10:45 AM
Costus deistelii
Page 186
K.Schum.
Zingiberaceae
Habitat
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).
Phenology
Description
Data sources
Guild: np
Life form: large herb
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
spp
186
Open (n)
C.d.
29
32 (19)
All
46333
37
FWTA, De Rouw (1991)
Distribution
Ghana
forest
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
14
38
41
34
23
69
24
52
17
39
37
29
24
10
69
25
36
13
39
11/11/03
10:45 AM
Page 187
Crossostemma laurifolium
Planch. ex Benth.
Passifloraceae
Description
Regeneration
Guild: pi
Life form: large woody climber
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.

Phenology
Distribution
dIvoire, Ghana
Forest type: montane evergreen forest, wet
evergreen forest, moist evergreen forest, secondary
forest (herbarium). In Cte dIvoire, 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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.l.
63
50 (46)
48
22
31
36
10
16
67
17
25
19
H
52
All
46333
37
39
37
29
24
10
69
25
36
13
39
187
11/11/03
10:45 AM
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
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.
Data sources
Distribution
Upper Guinea: Sierra Leone (FWTA), Liberia, Cte
dIvoire (herbarium)
Distribution type: Continuous distribution,
range is 871 km
Forest type: evergreen forests, riverine forest,
secondary forests
Habitat
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
Open (n)
C.g.
25
0 (14)
All
46333
37
Altitude
River
Coast
>500m
<2km
<5km
VW
72
52
28
16
96
68
20
39
37
29
24
10
69
25
36
13
39
Soil CMK
Soil WHC
11/11/03
10:47 AM
Page 189
Cussonia bancoensis
Aubrv. & 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
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
Other: it has a thick bark with deep fissures.
Regeneration
Phenology
Distribution
Continent: Nigeria (herbarium), Cameroon
(uncertain record, according to Bamps, 1974)
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 Aubrville (1959) and De Koning (1983)

to have been planted in Banco, Cte dIvoire 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
Open (n)
FWTA, Aubrville (1959), De la Mensbruge (1966),

Bamps (1974), De Koning (1983), Hawthorne
(1995a), IUCN Red List (2000), Kasparek (2000),
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.b.
12
29 (7)
58
33
41
25
17
25
58
17
H
58
All
46333
37
39
37
29
24
10
69
25
36
13
39
189
11/11/03
10:47 AM
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

Upper Guinea: Liberia, Cte dIvoire, Ghana.
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)
November (Liberia, Voorhoeve 1965); fruiting period
from December to January (Hawthorne 1995a)
Habitat
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 Cte dIvoire, 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
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
Open (n)
Regeneration
Uses

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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.a.
23
8 (13)
61
56
39
74
22
17
74
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:47 AM
Page 191
Cynometra leonensis
Hutch. & Dalziel
Leguminosae-Caes.
Description
Regeneration
Guild: np
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.

type (cf. Voorhoeve 1965). Seedlings are common
in the forest (Voorhoeve 1965), but experience a
high mortality in dense shade (Savill & Fox 1967).
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
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
Open (n)
Voorhoeve (1965), Savill & Fox (1967), Hawthorne

& Jongkind (2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.l.
14
75 (4)
43
93
86
14
86
All
46333
37
39
37
29
24
10
69
25
36
13
39
191
11/11/03
10:47 AM
Page 192
Cyrtorchis hamata
Schltr.
Orchidaceae
Distribution
Upper Guinea: Cte dIvoire, Ghana, Togo
Forest type: unknown
Habitat
Natural habitat unknown. Only few records from
gardens or mango trees in town (herbarium).
Phenology
Dispersal: by wind
Description
Guild: u
Life form: epiphytic herb
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
Open (n)
Data sources
FWTA
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
C.h.
- (0)
11
77
22
33
67
11
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:47 AM
Page 193
Dactyladenia dinklagei
(Engl.) Prance & F.White
Chrysobalanaceae
Description
Phenology
Guild: sb
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.
Data sources
Distribution

Habitat
It occurs in the understorey, often close to water
(e.g. riverbanks) (Hawthorne & Jongkind 2004)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.d.
50 (2)
33
22
33
22
33
11
22
56
22
22
11
H
44
All
46333
37
39
37
29
24
10
69
25
36
13
39
193
11/11/03
10:47 AM
Page 194
Dactyladenia smeathmannii
(Baill.) Prance & F.White
Chrysobalanaceae
Habitat
Found along secondary roads, riverbanks, and the
sea-side (herbarium, Hawthorne & Jongkind 2004).
Description
Phenology
Guild: u
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
Open (n)
Distribution
Distribution type: continuous, widespread,
Data sources
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
D.s.
50 (2)
50
17
83
83
17
All
46333
37
39
37
29
24
10
69
25
L
83
36
17
13
39
11/11/03
10:47 AM
Page 195
Dactyladenia whytei
(Stapf ) Prance & White
Chrysobalanaceae
Description
Phenology
Guild: np
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
Other: it starts as a shrub. The twigs are
pubescent.
Data sources
Distribution
Upper Guinea: Guinea, Sierra Leone, Liberia
(herbarium)
Forest type: rainforest, coastal forest, secondary
forest
Habitat
Species occurrence increases very sharply with
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.w.
32
67 (12)
44
13
89
91
69
All
46333
37
39
37
29
24
10
69
25
36
13
39
195
11/11/03
10:47 AM
Page 196
Deinbollia voltensis
Hutch. ex Burtt Davy & Hoyle (nom. nud.)
Sapindaceae
Description
Phenology
Guild: u
Life form: shrub
leaflets, elliptic to oblanceolate, microphyll (2-3.2 x
6.5-9 cm), entire, herbaceous, winged rachis
Flower: small, white
Fruit: fleshy, globose (0.8 cm in diameter), reddish
orange
Distribution
Data sources

Upper Guinea: Ghana
in 1 30 cell. It has only been recorded near the
Volta River and is possibly extinct.
Forest type: gallery forest
Habitat
Found on seasonally flooded grasslands (Hawthorne
& Jongkind 2004).
spp
196
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.v.
- (0)
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:47 AM
Page 197
Dennettia tripetala
Baker f.
Annonaceae
Description
Phenology
Guild: sb
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)

Distribution
Uses
Continent: Nigeria, Cameroon

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 Cte dIvoire, 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.
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.t.
16
27 (11)
75
19
75
25
25
19
H
56
All
46333
37
39
37
29
24
10
69
25
36
13
39
197
11/11/03
10:47 AM
Page 198
Dialium aubrevillei
Pellegr.
Leguminosae-Caes.
Regeneration
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).
Phenology
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
Ghana
semi-deciduous forest, secondary forest. In Ghana,
the species is most common in evergreen forests
Description
Guild: sb
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
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.a.
47
10 (31)
21
11
71
10
68
30
43
H
45
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:48 AM
Page 199
Diaphananthe suborbicularis
Summerh.
Orchidaceae
Description
Data sources
Guild: u
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)
Distribution
Upper Guinea: Ghana
in 1 30 cell. Only known from the type specimen
Westwood 158 from the Togo Plateau (Kpandu to
Wurupong)
Habitat
Natural habitat unknown. Only one record on a
cocoa tree (herbarium).
Phenology
Dispersal: by wind
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.s.
- (0)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
199
11/11/03
10:48 AM
Page 200
Dichapetalum albidum
A.Chev. ex Pellegr.
Dichapetalaceae
Description
Habitat
Guild: np
Max. diameter: 5 cm (Breteler 1973)
(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
type (cf. Breteler 1973).
Phenology
Dispersal: most probably by birds
Distribution
dIvoire (Breteler 1973), Ghana (herbarium)
Forest type: montane forest, rainforest, semideciduous forest, gallery forest, secondary forest
Data sources
Breteler (1973), Hall & Swaine (1981)
spp
200
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.a.
40
37 (19)
20
55
20
41
38
75
25
65
H
15
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:48 AM
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
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
type (cf. Breteler 1973) and regenerates in shade
(Hall & Swaine 1981) but occurs also in thickets
and secondary forest (herbarium).
Phenology
Timing: flowering is not clearly seasonal (Hall &
Swaine 1981)
Distribution
Continent: Benin, Nigeria, Gabon, Democratic
Republic of Congo, Congo (Brazzaville)
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
Open (n)
FWTA, Breteler (1973), Nwude et al. (1977), Hall &

Swaine (1981), Hawthorne & Jongkind (2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.b.
86 (7)
11
56
100
78
22
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
201
11/11/03
10:48 AM
Page 202
Dichapetalum dictyospermum
Breteler
Dichapetalaceae
Description
Regeneration
Guild: np
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.

type (cf. De Koning 1983) and can regenerate
abundantly (herbarium).
Phenology
Timing: flowering period from July to March;
fruiting period from October to April (De Koning
1983)
Distribution
Upper Guinea: Cte dIvoire (herbarium, Breteler
1978)
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.d.
53
38 (24)
92
100
98
98
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:48 AM
Page 203
Dichapetalum filicaule
Breteler
Dichapetalaceae
Description
Guild: np
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.
Phenology
Dispersal: by animals (Breteler 1973)
December and May to June; fruiting period from
December to January (De Koning 1983)
Distribution
spp
Continent: Upper Guinea endemic (Breteler 1978)

Breteler 1978)
in 11 30 cells
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).
Open (n)
Data sources
Breteler (1973, 1978), Hall & Swaine (1981), De
Koning (1983)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.f.
63
77 (31)
63
10
40
57
27
65
25
59
All
46333
37
39
37
29
24
10
69
25
36
13
39
203
11/11/03
10:48 AM
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
Regeneration
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
Max height: 100 m long (herbarium)
Max. diameter: 12 cm (herbarium, Breteler 1982)
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
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.
Distribution
Forest type: wet evergreen, moist evergreen,
riverine forest, moist semi-deciduous forest, secondary
spp
204
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
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
D.t.
142
62 (63)
45
11
47
20
25
69
25
42
H
43
All
46333
37
39
37
29
24
10
69
25
36
13
39
Data sources
Breteler (1973, 1982), Hall & Swaine (1981)
11/11/03
10:48 AM
Page 205
Didelotia engleri
Dinkl. & Harms
Leguminosae-Caes.
Description
Phenology
Guild: u
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
Data sources
FWTA, Oldeman (1964), Savill & Fox (1967), Lock
Distribution
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
D.e.
0 (1)
50
50
50
50
All
46333
37
39
37
29
24
10
L
0
6
Soil WHC
M
100
50
69
25
36
13
39
205
11/11/03
10:49 AM
Didelotia idae
Page 206
Oldeman, de Wit & J.Lonard
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.

type (cf. Voorhoeve 1965). Seedlings are prolific
after a seed year, and seem capable of
withstanding the dense shade (Savill & Fox 1967).
Phenology
Timing: flowering does not occur annually (Savill &
Fox 1967); fruiting period from October to
November (Liberia, Voorhoeve 1965)
Distribution
Ghana
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
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
Open (n)
Habitat
Uses

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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.i.
18
8 (13)
61
39
17
44
89
11
72
22
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
10:49 AM
Page 207
Diospyros chevalieri
De Wild.
Ebenaceae
Description
Regeneration
Guild: sb
Leaf: opposite, simple, oblong, mesophyll (6-11 x
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.
Phenology
Distribution
Ghana
forest, coastal forest, secondary forest. In Ghana,
most abundant in wet evergreen forests (Hall &
Swaine 1981).
Data sources
FWTA, White (1978), Hall & Swaine (1981),
Habitat
(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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
D.c.
79
28 (53)
35
71
11
19
68
30
61
38
All
46333
37
39
37
29
24
10
69
25
36
13
39
207
11/11/03
10:49 AM
Diospyros cooperi
Page 208
(Hutch. & Dalziel) F.White
Ebenaceae
Description
Regeneration
Guild: sb
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.
Phenology
Distribution
Ghana
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
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.c.
35
19 (21)
34
43
18
35
83
14
83
H
11
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:50 AM
Page 209
Diospyros liberiensis
Ebenaceae
Description
Phenology
Guild: pi
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
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
Distribution
FWTA, White (1978), Hawthorne & Jongkind

(2004)
Data sources

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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.l.
50 (4)
33
50
50
34
17
100
67
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
209
11/11/03
11:50 AM
Diospyros vignei
Page 210
F.White
Ebenaceae
Habitat
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
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.
Phenology
Distribution

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)

spp
210
Open (n)
Data sources
FWTA, White (1963, 1978), Hall & Swaine (1981),
Hawthorne (1995a)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.v.
37
34 (35)
24
19
56
24
65
30
35
57
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:50 AM
Page 211
Dissomeria crenata
Hook.f. ex Benth.
Flacourtiaceae
Description
Phenology
Guild: u
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)
Distribution
Data sources
Continent: Nigeria
Upper Guinea: Guinea, Sierra Leone, Cte dIvoire,
Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.c.
30
40 (5)
43
74
10
63
30
47
13
17
All
46333
37
39
37
29
24
10
69
25
36
13
39
211
11/11/03
11:50 AM
Page 212
Dorstenia astyanactis
Ak Assi
Moraceae
Description
Phenology
Guild: u
Life form: perennial, non-woody, succulent epiphyte
Max. height: 1.5 m (Ak Assi 1967)
Max. diameter: 1 cm (Ak 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
Other: it is a pendulous epiphyte.
Data sources
Distribution
FWTA, Ak Assi (1967), Hawthorne & Jongkind

(2004)

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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.a.
- (0)
100
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:50 AM
Page 213
Dorstenia embergeri
Mangenot
Moraceae
Description
Phenology
Guild: u
Life form: perennial herb
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.
Data sources
Mangenot (1957), De Koning (1983), Hawthorne &
Jongkind (2004)
Distribution
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.e.
12
0 (6)
75
17
75
17
58
25
17
67
All
46333
37
39
37
29
24
10
69
25
36
13
39
213
11/11/03
11:50 AM
Page 214
Dorstenia turbinata
Engl.
Moraceae
Description
Phenology
Guild: sb
Life form: small shrub or small tree
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.
Data sources
Distribution
FWTA, Hawthorne (1996), Hawthorne & Jongkind

(2004)

Gabon, Democratic Republic of Congo
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
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
D.t.
61
12 (42)
41
41
24
32
72
26
46
H
44
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:50 AM
Page 215
Dracaena calocephala
Bos
Dracaenaceae
Description
Phenology
Guild: sb
Life form: rhizotomatic shrub
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.
Timing: it appears not to have a fixed flowering period
Distribution
Data sources

Upper Guinea: Guinea, Liberia
Forest type: wet forest, secondary forest
Bos (1984)
Habitat
It occurs terrestrial in forest undergrowth,
presumably in undisturbed forests
(Bos, pers. comm.).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.c.
0 (3)
38
50
63
38
75
25
75
All
46333
37
39
37
29
24
10
69
25
36
13
39
215
11/11/03
11:50 AM
Page 216
Dracaena cristula
W.Bull
Dracaenaceae
Habitat
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).
Regeneration
Presumably by seed only.
Phenology
Timing: it does not appear to have a fixed
flowering period
Description
Guild: sb
Life form: unbranched pigmy tree
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
Open (n)
Distribution
dIvoire, Ghana (herbarium, Bos 1984)
swamp forest, secondary forest
Data sources
Bos (1984)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.c.
72
23 (35)
15
44
29
47
12
68
29
39
H
35
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:50 AM
Page 217
Dracaena ovata
Ker-Gawl.
Dracaenaceae
Description
Guild: sb
Life form: shrub
Max. diameter: less than 2 cm
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.
Distribution
Continent: Nigeria, Cameroon (Bos 1984)
Ghana (herbarium, Bos 1984)
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
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
frequently found near streams, rivers, or lakes
(herbarium). On sandy, clayish, rocky, and lateritic
soils (herbarium)
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
D.o.
82
42 (45)
39
15
54
20
18
56
37
28
H
5
All
46333
37
39
37
29
24
10
69
25
36
13
39
217
11/11/03
11:50 AM
Page 218
Dracaena phrynioides
Hook.
Dracaenaceae
Habitat
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
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.
Timing: it appears not to have a fixed flowering
period
Open (n)
Distribution
Continent: Nigeria (Bos 1984), Cameroon,
Equatorial Guinea (herbarium), Fernando Po (Bos,
1984), Gabon (herbarium)
(herbarium)
Upper Guinea the species is present in 21 30 cells,
distribution range is 2468 km
forest, moist semi-deciduous forest, dry semideciduous forest, riverine forest
Data sources
FWTA, Hall & Swaine (1981), Bos (1984)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.p.
30
7 (15)
10
33
10
73
17
37
60
20
H
60
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:54 AM
Page 219
Dracaena praetermissa
Bos
Dracaenaceae
Description
Phenology
Guild: sb
Life form: small shrublet
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.
flowering period
Distribution
Data sources
Continent: Upper Guinea endemic to the Cavally

river basin in SW Cte dIvoire
Upper Guinea: Cte dIvoire, presumably also on
the Liberian bank
3 30 cells, distribution range is 194 km.
forest
Bos (1984), Hawthorne & Jongkind (2004)
Habitat
It is found in deep shade.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.p.
29
20 (5)
97
86
86
All
46333
37
39
37
29
24
10
69
25
36
13
39
219
11/11/03
11:54 AM
Dracaena scabra
Page 220
Bos
Dracaenaceae
Description
Guild: np
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
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
Open (n)
Phenology
Timing: it appears not to have a fixed flowering
period
Distribution
Upper Guinea: Cte dIvoire (herbarium, Bos 1984)
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
D.s.
13
40 (10)
46
93
38
54
85
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:54 AM
Page 221
Ehretia trachyphylla
C.H.Wright
Boraginaceae
Description
Phenology
Guild: pi
Leaf: alternate, simple, elliptic, mesophyll (4-8 x 820 cm), entire, coriaceous, pubescent at lower
surface
Fruit: fleshy, narrowly obovoid (0.6 x 1 cm), white
to greyish; 4 seeds
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)
Distribution
Data sources

Forest type: evergreen forest, semi-deciduous
evergreen forest, secondary forest
FWTA, Hall & Swaine (1981),

De Koning (1983), Hawthorne (1990),
Habitat
It occurs in mature and secondary forests, and
sometimes along river borders (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
E.t
17
11
(9) 0
41
47
35
18
41
53
24
H
59
All
46333
37
39
37
29
24
10
69
25
36
13
39
221
11/11/03
11:54 AM
Page 222
Elytraria ivorensis
Dokosi
Acanthaceae
Distribution
Forest type: wet evergreen forest, old secondary
forest (herbarium)
Habitat
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
Description
Guild: sb
Life form: perennial rosette-forming herb
Leaf: rosette, simple, obovate, microphyll (2-3 x 46 cm), entire, pubescent above, glabrous beneath
Seed: small (0.1 cm), brown
spp
222
Open (n)
Data sources
FWTA, Dokosi (1970), Hawthorne & Jongkind (2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
E.i.
11
36 (11)
27
72
27
82
64
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:54 AM
Page 223
Elytraria maritima
J.K.Morton
Acanthaceae
Description
Phenology
Guild: pi
Life form: small rhizomatous annual herb
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)
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
E.m.
15
67 (9)
67
14
80
40
47
13
20
53
All
46333
37
39
37
29
24
10
69
25
36
13
39
223
11/11/03
11:54 AM
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)
Forest type: riverine forest, gallery forest in
savanna
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)
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
red with a sticky pulp; 1 seed
spp
224
Open (n)
Data sources
FWTA, Polhill & Wiens (1998)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
E.g.
- (0)
25
50
25
75
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:54 AM
Page 225
Englerina parviflora
(Tiegh.) Balle
Loranthaceae
Description
Phenology
Guild: pi
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
red or purple with a sticky pulp; 1 seed
Dispersal: by birds
Timing: flowering period from December to April
(Polhill & Wiens 1998)
Distribution
Continent: Upper Guinea endemic (Polhill & Wiens
1998)
Leone, Liberia, Cte dIvoire (Polhill & Wiens 1998),
Ghana (herbarium)
Data sources
Habitat
Data unavailable.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
E.p.
22
- (0)
23
41
59
28
10
45
55
36
All
46333
37
39
37
29
24
10
69
25
36
13
39
225
11/11/03
11:54 AM
Page 226
Euadenia eminens
Hook.f.
Capparaceae
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
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
Deciduousness: deciduous (herbarium)
Habitat
Distribution
spp
Phenology

Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
E.e
14
14 (7)
57
14
14
43
29
14
50
36
21
H
57
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:54 AM
Page 227
Ficus ottoniifolia
(Miq.) Miq. ssp.
multinervia
C.C.Berg
Moraceae
Description
Phenology
Guild: pi
Life form: shrub or tree, starts epiphytic
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
Other: the slash exudes a watery, whitish latex.
Dispersal: probably by animals (monkeys, birds,

bats)
Distribution
Data sources

Upper Guinea: Senegal, Gambia, Guinea-Bissau,
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
F.o.
18
33 (3)
44
33
51
22
11
11
56
33
39
11
H
28
All
46333
37
39
37
29
24
10
69
25
36
13
39
227
11/11/03
11:54 AM
Ficus scott-elliotii
Page 228
Mildbr. & Burrett.
Moraceae
Description
Phenology
Guild: u
Life form: tree, starts epiphytic
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
Other: the slash exudes a white latex.
Dispersal: probably by animals (monkeys, birds,

bats)
Distribution
Data sources

Sierra Leone, Cte dIvoire, Ghana
Forest type: moist forest, riverine forest
Habitat
Usually, found along rivers in moist forests
(herbarium).
spp
228
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
F.s.
14
25 (4)
50
21
72
14
43
57
36
29
14
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:57 AM
Page 229
Garcinia elliotii
Engl.
Guttiferae
Description
Phenology
Guild: u
Life form: shrub
Leaf: opposite, simple, oblong to oblong-elliptic,
notophyll (3-4 x 10-12 cm), entire, coriaceous,
glabrous; distinctly petiolate
Inflorescence: axillary, fascicle
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

Upper Guinea: Guinea Bissau, Sierra Leone
Habitat
It is sometimes found by streams (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.e.
- (0)
100
50
51
100
50
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
229
11/11/03
11:57 AM
Gardenia nitida
Page 230
Hook.
Rubiaceae
Distribution
Uses
Continent: Nigeria, Cameroon (herbarium)

Leone, Cte dIvoire, Ghana (herbarium)
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
Habitat
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.
Phenology
Description
Guild: sb
Leaf: opposite, simple, elliptic, mesophyll (3-8.5 x
7-25 cm), entire, becoming dentate towards the
apex, glabrous; interpetiolar stipules
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
Other: it has pale corky twigs, usually forking into
trees.
spp
230
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.n.
37
20 (15)
30
19
65
17
19
62
35
16
35
41
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:57 AM
Page 231
Gilbertiodendron bilineatum
(Hutch. & Dalziel) J.Lonard
Leguminosae-Caes.
Description
Phenology
Guild: u
Life form: medium-sized to large tree
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
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
Data sources
Distribution
FWTA, Savill & Fox (1967), Lock (1989), Hawthorne

(1995a), IUCN Red List (2000), Hawthorne &
Jongkind (2004)

Ghana
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.b.
17
0 (4)
29
12
30
30
30
65
35
41
H
53
All
46333
37
39
37
29
24
10
69
25
36
13
39
231
11/11/03
11:57 AM
Page 232
Gilbertiodendron limba
(Scott-Elliot) J.Lonard
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.
Dispersal: animals (e.g. monkeys) might eat the
seeds (De Koning 1983, Hawthorne 1995a),
otherwise short-distance explosive dehiscence
(Aubrville 1959)
Timing: flowering period from February to June (De
Koning 1983); fruiting period from July to
September (Ghana, Taylor 1960)
Distribution
Upper Guinea: Guinea, Sierra Leona, Liberia, Cte
dIvoire, Ghana
Forest type: evergreen forest, swamp forest,
riverine forest, secondary forest
Habitat
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
Max diameter: 20 cm (herbarium)
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
Open (n)
Data sources
Aubrville (1959), Taylor (1960), Voorhoeve (1965),
De Koning (1983), Alexander (1989), Lock (1989),
Hawthorne (1995a)
Regeneration
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.l.
36
0 (19)
17
53
44
39
47
50
22
56
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:57 AM
Page 233
Gilbertiodendron obliquum
(Stapf ) J.Lonard
Leguminosae-Caes.
Description
Guild: u
Life form: shrub
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
in 1 30 cell. An extremely rare species; only one
record known from A. Whyte found at Sinoe Basin.
Data sources
FWTA, Lock (1989)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.o.
- (0)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
233
11/11/03
11:57 AM
Page 234
Aubrv. & Pellegr.
Leguminosae-Caes.
Description
Data sources
Guild: u
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)
Fruit: dry dehiscent (pod), green with brown
indumentum
Aubrville (1959), IUCN Red List (2000), Hawthorne

& Jongkind (2004)
Distribution

species (Vulnerable). The species has a very
restricted distribution in southwest Cte dIvoire.
Habitat
It has been observed along roadsides. Two out of
five records are from trees occurring near water
(herbarium).
Phenology
spp
234
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.r.
0 (5)
50
100
83
17
50
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:57 AM
Page 235
Globimetula assiana
(Balle) Wiens & Polhill
Loranthaceae
Description
Phenology
Guild: pi
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
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

1998)
dIvoire (Polhill & Wiens 1998)
Forest type: montane forest
Polhill & Wiens (1998)
Habitat
It occurs between 0-1200 m altitude (Polhill & Wiens
1998).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.a.
25
- (0)
76
60
12
12
68
24
76
12
H
8
All
46333
37
39
37
29
24
10
69
25
36
13
39
235
11/11/03
11:57 AM
Page 236
Globimetula cupulata
(DC.) Danser
Loranthaceae
Description
Phenology
Guild: pi
Leaf: opposite, simple, ovate to broadly elliptic,
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)
Other: the twigs are flattened.
Dispersal: by birds
Timing: flowering period from December to May
Data sources
Distribution
1998)
Upper Guinea: Senegal, Guinea Bissau, Guinea,
Sierra Leone (Polhill & Wiens 1998)
Forest type: bushes on coastal dunes
Habitat
Commonly found on Parinari trees, but also other
trees (Polhill & Wiens 1998).
spp
236
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.c.
22
100 (3)
14
64
41
50
18
23
23
77
59
H
18
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
11:57 AM
Page 237
Guaduella macrostachys
(K.Schum.) Pilger
Gramineae
Description
Guild: sb
Life form: perennial herb (dwarf bamboo)
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
Other: it is violet at the base.
Distribution
Continent: Nigeria, Cameroon, Democratic

Republic of Congo (herbarium)
Upper Guinea: Ghana (herbarium, Hall & Swaine
1981)
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
Data sources
Hall & Swaine (1981), Van der Zon (1992)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.m
0 (3)
66
33
33
67
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
237
11/11/03
11:57 AM
Page 238
Guaduella oblonga
Hutch. ex W.D.Clayton
Gramineae
Description
Phenology
Guild: sb
Max. height: 1.2 m (Van der Zon 1992)
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
Data sources
FWTA, De Koning (1983), Van der Zon (1992),
Poilecot (1995)
Distribution
Continent: Cameroon, Equatorial Guinea

(herbarium), Gabon (Van der Zon 1992)
dIvoire (herbarium)
Habitat
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
G.o.
42
31 (13)
10
48
23
31
45
86
12
69
H
12
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:01 PM
Page 239
Guibourtia dinklagei
(Harms) J.Lonard
Leguminosae-Caes.
Description
Guild: u
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)
Fruit: dry indehiscent
Other: it has buttresses up to 1.5 m high.
Distribution
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),
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.d.
67 (3)
25
25
50
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
239
11/11/03
12:01 PM
Page 240
Gymnosiphon longistylus
(Benth.) Hutch.
Burmanniaceae
Distribution
Continent: Nigeria, Cameroon, Gabon
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).
Phenology
spp
240
Description
Data sources
Guild: sb
Life form: saprophytic herb
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
FWTA
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.l.
14
0 (12)
14
43
28
35
35
71
29
43
H
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:01 PM
Page 241
Gymnostemon zaizou
Aubrv. & Pellegr.
Simaroubaceae
Description
Phenology
Guild: sb
Leaf: alternate, imparipinnately compound; 13-25
subopposite leaflets, oblong, notophyll (2-4.5 x
8-13.5 cm), entire, coriaceous, glabrous
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

30 cells, distribution range is 201 km, Red List
species (Vulnerable). Recorded only in southwest
Cte dIvoire
Forest type: moist evergreen forest
Data sources
De Rouw (1991), IUCN Red List (2000), Kasparek
(2000)
Habitat
Data unavailable.
Regeneration
type (cf. de la Mensbruge 1966), and regenerates
in undisturbed, primary forests (De Rouw 1991).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
G.z.
0 (3)
43
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
241
11/11/03
12:01 PM
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
Description
Guild: u
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
Open (n)
Distribution
Continent: Gabon
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
>500m
<2km
<5km
VW
Soil CMK
L
75
17
17
67
25
36
13
39
H.c.
12
0 (2)
75
100
17
All
46333
37
39
37
29
24
10
Soil WHC
69
11/11/03
12:01 PM
Page 243
Heritiera utilis
Sprague
Sterculiaceae
Description
Guild: np
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
Ghana (herbarium)
semi-deciduous forest, gallery forest, secondary
forest. In Ghana, mostly found in wet evergreen
Habitat
(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
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 Cte dIvoire (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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
H.u.
62
11 (38)
44
50
24
26
74
23
45
48
All
46333
37
39
37
29
24
10
69
25
36
13
39
243
11/11/03
12:01 PM
Hibiscus whytei
Page 244
Stapf
Malvaceae
Description
Data sources
Guild: u
Life form: tall herb
Leaf: alternate, simple, ovate to triangular (3-9 cm
long), irregularly dentate
Inflorescence: solitary
Flower: large; corolla yellow with purple centre
FWTA
Distribution
Upper Guinea: Guinea (FWTA), Liberia (herbarium),
Cte dIvoire (FWTA)
in 1 30 cell
Habitat
Data unavailable.
Phenology
spp
244
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.w.
- (0)
100
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:01 PM
Page 245
Hippocratea vignei
Hoyle
Celastraceae
Description
Phenology
Guild: np
Life form: large irregularly winding woody climber
Leaf: opposite, simple, oblong, mesophyll (8 x 12
cm), dentate, coriaceous
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
Hall (1962), Hawthorne (1995a), Hawthorne

& Jongkind (2004)
Continent: Nigeria
range is 1384 km
Forest type: rainforest, semi-deciduous forest,
savanna, secondary vegetation
Habitat
It is found along roads and clearings, but also in
mature forest (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
Soil CMK
Soil WHC
VW
H.v.
11
22 (9)
27
27
73
55
45
45
45
All
46333
37
39
37
29
24
10
69
25
36
13
39
245
11/11/03
12:01 PM
Hugonia rufipilis
Page 246
Linaceae
Description
Phenology
Guild: np
Life form: large woody liana, climbing with hooks
11-32 cm), serrate, herbaceous, pubescent; nerves
pale brown and hairy
(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.
Ghana (herbarium)
FWTA, Badr (1971), Hall & Swaine (1981), De

Koning (1983)
Habitat
It is often found near rivers.
spp
246
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.r.
11
29 (7)
55
18
55
36
100
36
64
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:01 PM
Page 247
Hunteria ghanensis
J.B.Hall & Leeuwenb.
Apocynaceae
Description
Guild: sb
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
Other: the bark is pale grey, shallowly fissured, but
dark brown on section. It exudes a scanty white
latex.
Distribution

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
Open (n)
Hall & Leeuwenberg (1979), Hall & Swaine (1981),

Hawthorne (1995a, 1996), Omino (1996), IUCN Red
List (2000)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.g.
10
20 (5)
60
20
60
30
10
80
20
10
H
80
All
46333
37
39
37
29
24
10
69
25
36
13
39
247
11/11/03
12:01 PM
Page 248
Hutchinsonia barbata
Robyns
Rubiaceae
spp
248
Description
Distribution
Guild: np
Life form: straggling shrub
Leaf: opposite, simple, ovate to lanceolate,
microphyll (1.5-3 x 4-8 cm), entire, herbaceous,
hairy; almost sessile; interpetiolar stipules
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
Other: occasionally with spines on the lower parts
of the stems. The twigs are covered with dense
curly hairs.

Upper Guinea: Guinea (Hawthorne & Jongkind
2004), Sierra Leone, Liberia, Cte dIvoire
(herbarium)
in 20 30 cells, distribution range is
842 km
Forest type: montane forest, moist evergreen
forest, coastal forest, swamp forest, secondary
forest
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.
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.b.
29
43 (21)
34
17
41
24
34
86
14
66
H
34
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:04 PM
Page 249
Hutchinsonia glabrescens
Robyns
Rubiaceae
Description
Guild: sb
Life form: straggling shrub
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
Other: it has spines.
Distribution
Habitat
Data unavailable.
Phenology
Data sources
FWTA
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.g.
0 (1)
33
33
33
33
33
33
100
67
All
46333
37
39
37
29
24
10
69
25
36
13
39
249
11/11/03
12:04 PM
Page 250
Robbr.
Rubiaceae
Description
Phenology
Guild: sb
Life form: shrub
Max. height: 0.6 m (Robbrecht 1977)
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
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
Habitat
It occurs in the shaded understorey (Robbrecht
1977).
spp
250
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.a.
- (0)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:05 PM
Page 251
Hymenostegia gracilipes
Hutch. & Dalziel
Leguminosae-Caes.
Description
Phenology
Guild: sb
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
Hawthorne (1995a), IUCN Red List (2000),
Kasparek (2000), Hawthorne & Jongkind
(2004)
Distribution
Upper Guinea: Cte dIvoire (one doubtful record),
Ghana
5 30 cells, distribution range is 257 km, Red List
species (Endangered). In Ghana, it is an uncommon
species (Hawthorne 1995a).
Habitat
It is mostly seen close to rivers in evergreen forest
(Hawthorne & Jongkind 2004). On sandy soils
(herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
H.g.
0 (3)
83
34
67
33
33
33
17
67
All
46333
37
39
37
29
24
10
69
25
36
13
39
251
11/11/03
12:05 PM
Page 252
Illigera vespertilio
(Benth.) Baker f.
Hernandiaceae
Description
Phenology
Guild: np
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
Flower: small; corolla pale pink
Fruit: dry indehiscent (3 cm long), 2 or 3 opposite
wings
Other: it climbs with a winding petiole.
Dispersal: probably by wind
Distribution
Data sources
Continent: Cameroon, Gabon, Democratic Republic

of Congo
Ghana
Distribution type: continental disjunct, widespread,
Forest type: rainforest, gallery forest, secondary
forest
Kubitzki (1969), De Koning (1983),

Habitat
Data unavailable.
spp
I.v.
All
252
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
11
29 (7)
18
72
18
18
27
55
18
18
64
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:05 PM
Page 253
Impatiens nzoana
A.Chev. ssp.
bennae
Jacq.-Fl.
Balsaminaceae
Description
Phenology
Guild: sb
Max. height: 0.6 m
Leaf: spirally arranged, simple, narrowly elliptic to
oblong, notophyll (1.8-4 x 6-17 cm), finely dentate,
glabrous
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.
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 Cte dIvoire. 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.
Habitat
I. nzoana grows in moist shaded places of riverine
forests and along watercourses.
Regeneration
Spreading by thin underground rhizomes.
spp
I.n.
All
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
0 (2)
50
50
100
83
17
66
17
46333
37
39
37
29
24
10
69
25
36
13
39
253
11/11/03
12:05 PM
Isolona cooperi
Page 254
Hutch. & Dalziel ex Coop. & Record
Annonaceae
Distribution
Data sources

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),

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
Description
Guild: sb
14-28 cm), entire, coriaceous, very shining and
dark-green above
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
39
38 (29)
31
13
79
16
41
54
23
72
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:05 PM
Page 255
Isonema smeathmannii
Roem. & Schult.
Apocynaceae
Description
Habitat
Guild: pi
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.

rainfall higher than 2000 mm/yr to reach an
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
Timing: flowering during the whole year (Van der
Ploeg 1983)
Distribution
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
145
81 (52)
49
35
25
28
39
33
46
21
14
27
H
21
46333
37
39
37
29
24
10
69
25
36
13
39
255
11/11/03
12:05 PM
Ixora hiernii
Page 256
Scott-Elliot
Rubiaceae
Description
Phenology
Guild: sb
Max. height: 7 m (De Block 1998)
mesophyll (3-8 x 8-21.5 cm), fairly coriaceous,
glabrous; petiole 0.3-1.5 cm long, glabrous; with
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, Cte dIvoire,
Ghana, Togo (herbarium, De Block 1998)
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
28
0 (1)
54
43
11
25
50
15
32
68
21
H
7
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:05 PM
Page 257
Ixora laxiflora
Sm.
Rubiaceae
Description
Habitat
Guild: pi
Max. height: 15 m
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
Other: I. laxiflora var. laxiflora differs from
I. laxiflora var. linderi in its long pedicels.

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, Cte dIvoire, 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 Cte dIvoire. 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.
Data sources
FWTA, De Koning (1983), Hawthorne (1995a), De
Block (1998)
spp
I.l.
All
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
112
39 (23)
40
46
29
24
36
17
51
33
24
14
H
22
46333
37
39
37
29
24
10
69
25
36
13
39
257
11/11/03
12:05 PM
Ixora liberiensis
Page 258
De Block
Rubiaceae
Distribution
Upper Guinea: Liberia (herbarium, De Block 1998)
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
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
Open (n)
Data sources
De Block (1998)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
H
I.l.
- (0)
50
50
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:07 PM
Page 259
Ixora tenuis
De Block
Rubiaceae
Description
Guild: sb
Life form: shrub
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
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
Distribution
Upper Guinea: Ghana (herbarium, De Block 1998)
30 cells, distribution range is 254 km. Known only
from Ghana, although Hawthorne & Jongkind (2004)
expect it to occur in Cte dIvoire as well.
evergreen forest
Phenology
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
10
0 (2)
10
30
90
10
10
90
30
46333
37
39
37
29
24
10
69
25
36
13
39
259
11/11/03
12:07 PM
Keetia bridsoniae
Page 260
Jongkind
Rubiaceae
Habitat
Sometimes found along roads or near riverbanks.
Phenology
Description
Guild: u
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
green; infructescence on stalks; 1-2 seeds
Other: the main stem is quadrangular.
Data sources
K
Distribution
spp
260
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
K.b
13
50 (8)
46
61
15
23
54
46
54
23
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:07 PM
Page 261
Keetia obovata
Jongkind
Rubiaceae
Description
Guild: u
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
Fruit: green; 1-2 seeds
Distribution
Habitat
Data unavailable.
Phenology
Data sources
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
K.o.
50(2)
50
50
50
100
50
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
261
11/11/03
12:07 PM
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
Phenology
Dispersal: by animals (monkeys)
Description
Guild: np
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.
spp
262
Open (n)
Distribution
Data sources

Forest type: rainforest, secondary forest, thicket.
In Ghana, only found in evergreen forest, especially
upland forests but also present in moist and wet
FWTA, Hall & Swaine (1981), Persoon et al. (1992),

Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
L.m.
71
29 (42)
51
51
35
10
73
23
24
62
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:07 PM
Page 263
Landolphia togolana
(Hallier f.) Pichon
Apocynaceae
Description
Habitat
Guild: np
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
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: Cte dIvoire, Ghana, Togo
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).
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
L.t.
29
50 (8)
34
24
76
20
14
48
38
21
14
H
38
All
46333
37
39
37
29
24
10
69
25
36
13
39
263
11/11/03
12:07 PM
Page 264
Lasiodiscus mannii
Hook.f.
Rhamnaceae
Description
Habitat
Guild: sb
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
Distribution
Continent: Guineo-Congolian wide: from Cte
dIvoire to Democratic Republic of Congo and
Tanzania (Hall & Swaine 1981). Central African
Republic, Cameroon, Equatorial Guinea, Sao Tom
and Principe, Gabon, Congo (Brazzaville),
Democratic Republic of Congo (herbarium)
Upper Guinea: Liberia, Cte dIvoire (herbarium)
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).
spp
L.m.
All
264
Open (n)
Data sources
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
0 (2)
75
25
25
25
25
100
50
25
25
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:07 PM
Page 265
Leptoderris miegei
Ak Assi & Mangenot
Leguminosae-Pap.
Description
Phenology
Guild: np
Max. height: 15 m long (Ak Assi & Mangenot
1975)
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, Ak Assi & Mangenot (1975), Hawthorne
(1995a), Lock (1989)
Distribution
Habitat
In Cte dIvoire, it is found in the lagoon region,
though not always near water (Ak Assi & Mangenot
1975). It is fairly common in Ghana in swamps of
evergreen forest zones (Hawthorne 1995a).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
L.m.
12
50 (6)
83
16
83
50
50
42
42
All
46333
37
39
37
29
24
10
69
25
36
13
39
265
11/11/03
12:07 PM
Page 266
Leucomphalos libericus
Breteler
Leguminosae-Pap.
Habitat
Description
Data unavailable.
Guild: u
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
Phenology
Distribution
Data sources

spp
266
Open (n)
FWTA, Breteler (1994b), Hawthorne & Jongkind

(2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
L.l.
100 (3)
50
50
50
100
50
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:07 PM
Page 267
Loesenera kalantha
Harms
Leguminosae-Caes.
Description
Phenology
Guild: u
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)
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
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
L.k.
10
50 (4)
20
20
40
20
20
50
100
60
H
0
All
46333
37
39
37
29
24
10
69
25
36
13
39
267
11/11/03
12:08 PM
Page 268
Macropodiella garrettii
(C.H.Wright) C.Cusset
Podostemaceae
Description
Data sources
Guild: u
Life form: aquatic herb
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
Upper Guinea: Sierra Leone, Cte dIvoire
Habitat
Growing submerged or at water level on rocks, near
waterfalls or in fast-flowing rivers.
Phenology
spp
268
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.g.
100 (1)
50
25
50
50
25
75
25
50
25
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:11 PM
Page 269
Maesa nuda
Hutch. & Dalziel
Myrsinaceae
Description
Phenology
Guild: u
Max. height: 5 m long, 5 m high (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, Cte dIvoire
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 (Cte dIvoire) to very degraded (Cameroon)
mountain forests (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.n.
25 (4)
20
60
40
40
60
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
269
11/11/03
12:11 PM
Page 270
Magnistipula cupheiflora
Mildbr. ssp.
leonensis
F.White
Chrysobalanaceae
Description
Data sources
Guild: u
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
Upper Guinea: Sierra Leone, Cte dIvoire
in 3 30 cells
Habitat
Data unavailable.
Phenology
spp
270
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.c.
0 (1)
33
33
66
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:11 PM
Page 271
Magnistipula zenkeri
Engl.
Chrysobalanaceae
Description
Phenology
Guild: u
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
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

Ghana
Guinea present in 5 30 cells, a rare species
(Hawthorne & Jongkind 2004)
Habitat
Found at the border of rivers and swamps (Vivien &
Faure 1985, Hawthorne & Jongkind 2004).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.z.
0 (1)
50
17
50
17
17
67
33
17
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
271
11/11/03
12:11 PM
Page 272
Malaxis melanotoessa
Summerh.
Orchidaceae
Description
Data sources
Guild: u
Life form: terrestrial herb
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
Seed: very small
FWTA, Cribb (1978), Summerhayes (1958)
Distribution
in 1 30 cell. It has been found only in the Gola
Forest of Liberia approx. 28 km south of Ba by
Bunting
Habitat
It has been found growing on humus on very wet
rocks (herbarium).
Phenology
Dispersal: by wind
spp
272
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.m
- (0)
100
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:11 PM
Page 273
Mangenotia eburnea
Pichon
Asclepiadaceae
Description
Regeneration
Guild: np
Leaf: opposite, simple, oblong, microphyll (1-2.3 x
2.5-7 cm), entire, herbaceous
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.
Phenology
Dispersal: by wind (Hall & Swaine 1981)
Distribution
Continent: Nigeria
Upper Guinea: Senegal, Guinea-Bissau, Sierra
Forest type: In Cte dIvoire 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
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.e.
31
30 (20)
16
16
42
26
10
22
19
45
35
23
35
H
32
All
46333
37
39
37
29
24
10
69
25
36
13
39
273
11/11/03
12:11 PM
Page 274
Manotes macrantha
(Gilg) Schellenb.
Connaraceae
Description
Distribution
Guild: np
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
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
Phenology
Data sources
FWTA, Jongkind (1989), Hawthorne & Jongkind
(2004)
spp
274
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.m
16
50 (10)
25
63
19
43
100
31
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:11 PM
Page 275
Mapania baldwinii
Nelmes
Cyperaceae
Description
Habitat
Guild: sb
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
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 Cte dIvoire, this species is especially
found on slopes (Lorougnon 1972). In Cte dIvoire,
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

1981)
(herbarium, Hall & Swaine 1981, Simpson 1992)
forest, secondary forest. In Ghana mostly in wet
evergreen forests (Hall & Swaine 1981). It is locally
common.
Pollination: probably pollinated by wind (Hall &
Swaine 1981) and insects (Simpson 1992)
Dispersal: not known (Hall & Swaine 1981)
Regeneration
It regenerates in the shade.
Data sources
Lorougnon (1972), Hall & Swaine (1981), Simpson
(1992)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.b.
43
13 (24)
42
77
19
63
35
49
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
275
11/11/03
12:11 PM
Page 276
Mapania ivorensis
(Raynal) Raynal
Cyperaceae
Description
Phenology
Guild: sb
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
Dispersal: pollination probably by insects (Simpson

1992)
Data sources
Lorougnon (1972), Simpson (1992)
Distribution
Upper Guinea: Liberia and Cte dIvoire
(herbarium, Simpson 1992)
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).
spp
276
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.i.
29
45 (11)
28
83
10
41
59
31
69
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:11 PM
Page 277
Mapania linderi
Hutch. ex Nelmes
Cyperaceae
Description
Phenology
Guild: sb
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
Distribution
dIvoire (herbarium, Simpson 1992)
Forest type: rainforest, riverine forest, secondary
forests. It is locally common.
Habitat
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.l.
38
30 (20)
21
37
42
45
13
76
24
58
H
32
All
46333
37
39
37
29
24
10
69
25
36
13
39
277
11/11/03
12:11 PM
Mapania minor
Page 278
(Nelmes) J.Raynal
Cyperaceae
Description
Phenology
Guild: sb
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
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
Fruit: dry indehiscent, obovate (0.1 x 0.1 cm),
black, exocarp hard and rugulose; 1 seed
Seed: small
Other: the rhizome grows vertical.
Data sources
Distribution
Upper Guinea: Liberia, Cte dIvoire (herbarium,
Simpson 1992)
Habitat
between 1500-2000 mm/yr (Chi2 test). It is found in
moist and shady places of the forest (Simpson
1992, herbarium).
spp
278
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.m
13
13 (8)
62
15
69
30
69
31
46
38
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:14 PM
Page 279
Mapania rhynchocarpa
Lorougnon & J.Raynal
Cyperaceae
Description
Phenology
Guild: sb
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
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
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).
Data sources
Distribution
Continent: Nigeria (herbarium, Simpson 1992)
(herbarium), Ghana (Simpson 1992)
Habitat
Found in moist and shady places of the forest.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.r.
- (0)
20
80
80
20
20
80
20
60
All
46333
37
39
37
29
24
10
69
25
36
13
39
279
11/11/03
12:15 PM
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
Ghana (herbarium)
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
Max diameter: 80 cm (Voorhoeve 1965)
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
Data sources
Fruit: fleshy
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.a.
21
11 (9)
33
53
10
33
81
19
71
29
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:15 PM
Page 281
Marantochloa cuspidata
(Rosc.) Milne-Redh.
Marantaceae
Description
Habitat
Data sources
Guild: pi
Life history: large perennial rhizomatous herb
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

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
Upper Guinea: Senegal (Dhetchuvi 1996), Guinea,
(herbarium)
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
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
M.c.
87
29 (52)
44
30
29
30
71
28
56
23
All
46333
37
39
37
29
24
10
69
25
36
13
39
281
11/11/03
12:15 PM
Page 282
Marattia odontosora
Christ
Marattiaceae
Description
Guild: u
Life form: large perennial herb (fern)
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
largest population in Upper Guinea, present in 5 30
cells in Upper Guinea
Habitat
It is common above 600 m (FWTA)
Phenology
Data unavailable.
Data sources
FWTA
spp
282
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.o.
- (0)
50
17
17
84
17
83
17
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:15 PM
Page 283
Maschalocephalus dinklagei
Gilg & K.Schum.
Rapateaceae
Description
Phenology
Guild: sb
Life form: rhizomatous herb
Leaf: rosette, simple, linear to lanceolate,
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
Data sources
FWTA
Distribution
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, Cte dIvoire).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.d.
15
29 (7)
27
47
26
20
73
27
67
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
283
11/11/03
12:15 PM
Page 284
Memecylon aylmeri
Hutch. & Dalziel
Melastomataceae
Description
Regeneration
Guild: sb
Life form: pigmy tree or shrub
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
Other: the branches are quadrangular.

type (cf. Jacques-Flix 1978).
Phenology
Distribution
dIvoire, Ghana
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-Flix (1978), Hall & Swaine (1981)
Habitat
Species occurrence increases at higher altitudes
(logistic regression analysis, Chi2 test) and with
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).
spp
284
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.a.
30
25 (12)
27
43
23
47
23
63
37
53
H
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:15 PM
Page 285
Mendoncia combretoides
(A.Chev.) Benoist
Acanthaceae
Description
Phenology
Guild: np
Life form: large woody winding climber
Leaf: opposite, simple, elliptic to ovate, notophyll
(3-7.5 x 6.5-12 cm), entire, coriaceous; hairy on the
nerves and the petiole
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

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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.c.
32
24 (21)
53
57
43
59
41
25
66
All
46333
37
39
37
29
24
10
69
25
36
13
39
285
11/11/03
12:15 PM
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)
Timing: flowering period from January to March,
later (till June) in Guinea and Senegal (Berg 1982)
Distribution
Upper Guinea: Senegal, Gambia, Guinea-Bissau,
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
(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
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).
Open (n)
Data sources
Voorhoeve (1965), Hall & Swaine (1981), Berg
(1982), De Koning (1983), UCN Red List (2000),
Regeneration
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
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.r.
48
39 (31)
46
10
31
26
25
17
52
40
29
10
H
42
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:15 PM
Page 287
Millettia leonensis
Hepper
Leguminosae-Pap.
Description
Phenology
Guild: u
Life form: small tree, sometimes climbing
opposite leaflets, ovate to elliptic, microphyll (2.53.5 x 5-9 cm), herbaceous, pubescent beneath,
rachis with appressed reddish to purplish hairs
Flower: calyx brown; corolla purple, petals on the
outside covered with purplish hairs
Fruit: dry dehiscent, flat, woody, young pods
densely pilose
Data sources
Distribution
FWTA, Lock (1989), Hawthorne & Jongkind (2004)

in 2 30 cells
Forest type: high forest
Habitat
One of the two known specimens was collected
along a rivers edge (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.l.
50 (2)
100
100
100
50
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
287
11/11/03
12:15 PM
Millettia liberica
Page 288
Jongkind
Leguminosae-Pap.
Distribution
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
opposite leaflets, elliptic, herbaceous, almost
glabrous beneath
Flower: calyx whitish with purple spots; corolla
violet
Fruit: dry dehiscent, flat, woody
Other: it has twigs with many small lenticels.
spp
288
Open (n)
Data sources
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.l.
67 (6)
100
100
83
17
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:18 PM
Page 289
Millettia pallens
Stapf
Leguminosae-Pap.
Description
Phenology
Guild: pi
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

dIvoire
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
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
M.p.
59
82 (33)
41
12
90
68
32
58
12
12
All
46333
37
39
37
29
24
10
69
25
36
13
39
289
11/11/03
12:18 PM
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
Phenology
Description
Guild: sb
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
Other: sometimes found as an irregularly winding
shrub in thickets. It has a quadrangular stem and
tendril forming branchlets.
spp
290
Open (n)
Distribution
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
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
M.w
27
47 (17)
78
19
18
63
19
70
11
15
11
H
59
All
46333
37
39
37
29
24
10
69
25
36
13
39
Data sources
Hall & Swaine (1981)
11/11/03
12:18 PM
Page 291
Monocyclanthus vignei
Keay
Annonaceae
Description
Phenology
Guild: sb
Max. diameter: 15 cm (Keay 1953)
Flower: medium-sized (1.4-1.6 cm long), corolla
yellow
Fruit: with 7 free carpels, hairy
Other: it has a straight stem. The bark is smooth
with some large lenticels.
Data sources
FWTA, Keay (1953), Achenbach & Frey (1992),
Distribution
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).
range is 1005 km, Red List species (Endangered)
Forest type: rainforest (herbarium)
Habitat
An understorey tree found in rainforests (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.v.
25 (4)
57
14
14
72
86
14
71
29
All
46333
37
39
37
29
24
10
69
25
36
13
39
291
11/11/03
12:18 PM
Page 292
Monosalpinx guillaumetii
N.Hall
Rubiaceae
Description
Phenology
Guild: u
Life form: shrub
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

forest
Hall (1968), Hawthorne & Jongkind (2004)
Habitat
Data unavailable.
spp
292
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.g.
33 (3)
33
83
17
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:18 PM
Page 293
Mostuea hymenocardioides
Hutch. & Dalziel
Gelsemiaceae
Description
Phenology
Guild: u
Life form: shrub
Leaf: opposite, simple, microphyll (0.8-2.4 x 1.5-5
cm)
Flower: corolla white to pink, red inside; unpleasant
scent
Fruit: dry dehiscent (capsule), medium-brown,
shining
Distribution
Data sources

Forest type: rainforest, gallery forest
FWTA
Habitat
Found in wet places, near rivers and
in flooded areas (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.h.
15
0 (1)
73
47
20
26
40
60
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
293
11/11/03
12:18 PM
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
(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.
spp
294
Open (n)
Distribution
dIvoire
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
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
M.a.
69
77 (35)
10
42
25
24
23
47
87
12
57
H
23
All
46333
37
39
37
29
24
10
69
25
36
13
39
Data sources
11/11/03
12:18 PM
Page 295
Mussaenda grandiflora
Benth.
Rubiaceae
Description
Guild: np
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
dIvoire
in 23 30 cells, distribution range is 725 km. It can
be locally common.
forest, thickets
Phenology
Habitat
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
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
36
23
39
91
82
11
24
10
69
25
36
13
39
M.g.
44
81 (26)
43
All
46333
37
39
37
29
Soil WHC
295
11/11/03
12:18 PM
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
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;
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.
spp
296
Open (n)
Distribution
Continent: Nigeria
forest, savanna, swamp forest, secondary forest,
thickets, coastal shrubland
Data sources
FWTA, De Koning (1983), Hawthorne & Jongkind
(2004)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.t.
74
41 (44)
50
62
35
12
46
42
74
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:18 PM
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
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
Phenology
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
dIvoire, Ghana
Forest type: evergreen (wet, moist, upland), semideciduous (moist, dry), dry forest, secondary forest
(herbarium). In Ghana, it is found mostly in moist
Data sources
FWTA, De Ruiter (1976), Hall & Swaine (1981),

De Koning (1983), De Rouw (1991), Swaine (1996),
Habitat
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
M.l.
79
44 (34)
48
12
37
33
19
63
32
25
H
63
All
46333
37
39
37
29
24
10
69
25
36
13
39
297
11/11/03
12:18 PM
Page 298
Napoleonaea heudelotii
A.Juss.
Lecythidaceae
Description
Data sources
Guild: u
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
Other: it has whorled branches.
FWTA, Liben (1971b), Hawthorne & Jongkind (2004)
Distribution
Upper Guinea: Guinea, Sierra Leone, Liberia
Forest type: high forest, secondary regrowth,
gallery forest
Habitat
Data unavailable.
Phenology
spp
298
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
N.h.
15
33 (3)
87
87
73
27
33
47
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:23 PM
Page 299
Neostenanthera hamata
(Benth.) Exell
Annonaceae
Description
Guild: sb
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
Other: The twigs are pubescent.
Distribution
dIvoire, Ghana
Forest type: rainforest, swamp forest (herbarium),
evergreen forest (Hawthorne & Jongkind 2004)
Phenology
Habitat
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
FWTA, Irvine (1961), IUCN Red List (2000),

spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
N.h.
39
40
(20) 5
69
26
52
20
85
10
56
31
All
46333
37
39
37
29
24
10
69
25
36
13
39
299
11/11/03
12:23 PM
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
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
(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
type (cf. Voorhoeve 1965).
Phenology
Dispersal: by wind
Timing: flowering period from August to
September; fruiting period from December to
January
Distribution
Ghana
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).
riverine forest, secondary forest
Habitat
300
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)

spp
Uses
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
N.a.
25
7 (15)
40
44
16
40
72
24
60
28
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:23 PM
Page 301
Newtonia elliotii
(Harms) Keay
Leguminosae-Mim.
Description
Phenology
Guild: u
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
Flower: corolla white; fragrant
Fruit: flat, somewhat curved (2 x 8 cm), smooth
Seed: winged
Dispersal: by wind
Distribution
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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
N.e.
0 (2)
67
11
89
100
56
44
All
46333
37
39
37
29
24
10
69
25
36
13
39
301
11/11/03
12:24 PM
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 Cte
dIvoire to Cameroon, with a different variety in the
Democratic Republic of Congo (Keay 1989,
Hawthorne 1995a)
Ghana (herbarium)
Data sources
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)
spp
302
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
O.a.
11
0 (3)
36
18
72
45
55
45
36
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:24 PM
Page 303
Oleandra ejurana
Adams
Davalliaceae
Description
Guild: u
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
1981)
Upper Guinea: Ghana (herbarium, Hall & Swaine
1981)
Forest type: dry forest, woodland
Habitat
On trees and in shady and very moist sheltered
places on steep rocks.
Phenology
Data sources
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
O.e.
0 (3)
25
75
25
75
25
75
H
25
All
46333
37
39
37
29
24
10
69
25
36
13
39
303
11/11/03
12:24 PM
Oncoba brevipes
Page 304
Stapf
Flacourtiaceae
Distribution
Data sources

Upper Guinea: Guinea (FWTA), Sierra Leone,
Liberia, Cte dIvoire (herbarium)
secondary forest, swamp forest
FWTA, Aubrville (1959), Sleumer (1974),

Habitat
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
Description
Guild: np
15-22 cm), entire, clustered at terminal axis,
coriaceous, dark green above and paler beneath
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
spp
304
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
O.b.
52
59 (27)
44
19
25
22
41
98
65
13
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:24 PM
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)
2004)
Leaf: alternate, simple, heart-shaped, mesophyll
(12-15 cm long), entire, glabrous above, pubescent
beneath
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),

Continent: Guinea wide: from Sierra Leone to

Cameroon (Hall & Swaine 1981). Nigeria,
Cameroon, Equatorial Guinea (herbarium)
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
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.g.
11
29 (7)
36
45
18
36
55
45
27
36
All
46333
37
39
37
29
24
10
69
25
36
13
39
305
11/11/03
12:24 PM
Page 306
Pararistolochia mannii
(Hook.f.) Keay
Aristolochiaceae
Distribution
Continent: Benin, Nigeria, Congo (Brazzaville)
Habitat
Found in dense and open forests, sometimes near
lagoons. On sandy soil (herbarium).
Phenology
Description
Guild: u
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
Data sources
FWTA, Poncy (1978), Hawthorne & Jongkind (2004)
spp
306
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.m.
12
50 (4)
42
17
42
33
25
42
17
42
50
H
33
All
46333 3
39
37
29
24
10
69
25
36
13
39
11/11/03
12:24 PM
Page 307
Pavetta akeassii
J.B.Hall
Rubiaceae
Description
Phenology
Guild: sb
Life form: shrub
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
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)

evergreen forest, dry forest, secondary forest
Habitat
Typically found in the forest understorey. On sandy
or clayish soils (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.a.
13
0 (12)
23
54
85
31
69
23
23
All
46333
37
39
37
29
24
10
69
25
36
13
39
307
11/11/03
12:24 PM
Page 308
Pavetta micheliana
J.G.Adam
Rubiaceae
Distribution
Upper Guinea: Guinea (Adam 1973), Liberia,
Cte dIvoire (herbarium)
Forest type: evergreen forest, secondary
regrowth
Habitat
Sometimes close to streams (herbarium).
Phenology
Description
Guild: u
Max. height: 9 m
Leaf: opposite, simple, elliptic to ovate, macrophyll
(9-11 x 25-27.5 cm), entire, coriaceous, glabrous;
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
Data sources
Adam (1973), Hawthorne & Jongkind (2004)
spp
308
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.m.
33 (3)
50
25
25
75
100
75
25
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:40 PM
Page 309
Penianthus patulinervis
Hutch. & Dalziel
Menispermaceae
Description
Regeneration
Guild: sb
Life form: shrub
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
Ghana
(herbarium)
Uses
The roots are chewed as an aphrodisiac
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)
spp
P.p.
All
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
88
27 (59)
57
48
40
75
25
16
H
80
46333
37
39
37
29
24
10
69
25
36
13
39
309
11/11/03
12:40 PM
Page 310
Phragmanthera vignei
Balle
Loranthaceae
Description
Phenology
Guild: pi
(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
Data sources
FWTA, Balle (1956), Polhill & Wiens (1998)
Distribution
1998)
dIvoire, Ghana (herbarium, Polhill & Wiens 1998)
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).
Habitat
It occurs from the lowlands up to 1600 m altitude
(Polhill & Wiens 1998).
spp
310
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.v.
17
0 (1)
65
65
12
36
59
24
71
18
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:40 PM
Page 311
Pierreodendron kerstingii
(Engl.) Little
Simaroubaceae
Description
Habitat
Guild: sb
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
Other: it has a rough bark in square scales, with a
strong almond scent.
Sometimes found on riverbanks.
Phenology
Data unavailable.
Data sources
FWTA, Aubrville (1959), Keay (1989),
Hawthorne (1995a), IUCN Red List
Distribution
(herbarium), Togo (Hawthorne 1995a)
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, Cte dIvoire, 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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.k.
0 (1)
50
38
13
38
38
13
50
13
38
13
38
All
46333
37
39
37
29
24
10
69
25
36
13
39
311
11/11/03
12:40 PM
Page 312
Piptostigma fugax
Annonaceae
Habitat
It is found in the understorey of both dense and
disturbed forests. Occasionally reported on sandy
soils (herbarium).
Regeneration
Phenology
Dispersal: probably dispersed by animals
Description
Data sources
Distribution
Guild: sb
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)

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
spp
312
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.f.
16
23 (13)
13
44
19
57
19
81
19
56
H
31
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:40 PM
Page 313
Platysepalum hirsutum
(Dunn) Hepper
Leguminosae-Pap.
Description
Guild: np
Life form: medium-sized woody climber
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
dIvoire, Ghana
forest, dry semi-deciduous forest, secondary forest.
In Ghana, it occurs predominantly in drier forest
types (Hall & Swaine 1981).
Data sources
Habitat

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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.h.
65
17 (29)
35
38
32
13
19
55
43
38
51
All
46333
37
39
37
29
24
10
69
25
36
13
39
313
11/11/03
12:40 PM
Page 314
Pleiocoryne fernandensis
(Hiern) S.Rauschert
Rubiaceae
Description
Habitat
Guild: np
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
type (cf. De Koning 1983).
Phenology
Distribution
Fernando Po, Gabon, Congo, Democratic Republic
of Congo (Kirkbride & Robbrecht 1984), Angola
(herbarium)
Upper Guinea: Guinea (Kirkbride & Robbrecht
1984), Sierra Leone, Liberia, Cte dIvoire, Ghana
(herbarium), Togo (Kirkbride & Robbrecht 1984)
forest, coastal forest, dry forest, swamp forest,
secondary forest, thickets
Data sources
De Koning (1983), Kirkbride & Robbrecht (1984),
spp
314
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.f.
26
45 (11)
35
15
19
30
31
20
31
31
38
19
23
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:40 PM
Page 315
Poecilocalyx stipulosa
(Hutch. & Dalziel) N. Hall
Rubiaceae
Description
Phenology
Guild: sb
Life form: shrub
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
Other: the young twigs are softly hairy.
Data sources
Distribution
FWTA, De Rouw (1991), Hawthorne & Jongkind

(2004)

Habitat
It is found in the forest understorey (herbarium, De
Rouw 1991).
spp
Open (n)
P.s.
0 (1)
All
46333
37
Altitude
River
Coast
>500m
<2km
<5km
VW
67
33
66
83
17
83
39
37
29
24
10
69
25
36
13
39
Soil CMK
Soil WHC
315
11/11/03
12:40 PM
Page 316
Polycephalium capitatum
(Baill.) Keay
Icacinaceae
Description
Phenology
Guild: pi
Max. height: 3 m long
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
Distribution
FWTA

dIvoire
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).
spp
316
Open (n)
Data sources
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
P.c.
25
67 (9)
48
36
20
44
80
12
44
36
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
12:40 PM
Page 317
Polystachya bancoensis
Burg
Orchidaceae
Description
Phenology
Guild: pi
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
November (De Koning 1983)
Data sources
FWTA, Arends et al. (1980), De Koning (1983)
Distribution
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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.b.
0 (4)
63
38
63
25
63
13
25
25
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
317
11/11/03
12:40 PM
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.
Dispersal: by wind
Timing: flowering period from February to March
(Johansson 1974)
Data sources
FWTA, Jaeger et al. (1968), Johansson (1974)
Distribution
dIvoire
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).
spp
318
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.d.
17
25 (4)
88
18
24
53
18
71
29
41
H
29
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:42 PM
Page 319
Polystachya pseudodisa
Kraenzl.
Orchidaceae
Description
Guild: u
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
Seed: very small
Other: it has a pseudobulb.
Distribution
Habitat
It grows on trees or rocky slopes (herbarium).
Phenology
Dispersal: by wind
Data sources
FWTA
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.p.
- (0)
38
63
25
76
50
50
13
H
38
All
46333
37
39
37
29
24
10
69
25
36
13
39
319
11/11/03
3:42 PM
Page 320
Premna grandifolia
A.Meeuse
Verbenaceae
Description
Data sources
Guild: u
Life form: shrub
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
Fruit: unknown
Seed: unknown
FWTA, IUCN Red List (2000), Hawthorne & Jongkind

(2004)
Distribution
Distribution type: present in 2 30 cells. It is only
known from southwest Cte dIvoire. A Red list
species (Vulnerable).
Habitat
Data unavailable.
Phenology
spp
320
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.g.
- (0)
50
50
50
50
50
50
H
0
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:42 PM
Page 321
Breteler
Acanthaceae
Description
Phenology
Guild: u
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
Other: it has grey-green stems.
Data sources
Distribution
Breteler (1994a, 1998), Hawthorne & Jongkind

(2004)

Upper Guinea: Liberia (Breteler 1994a, herbarium)
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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.l.
67 (3)
50
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
321
11/11/03
3:42 PM
Page 322
Pteleopsis habeensis
Aubrv. ex Keay
Combretaceae
Description
Guild: pi
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
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 seasons 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),
Phenology
Deciduousness: deciduous (Geerling 1984)
Dispersal: by wind
Distribution
Continent: Mali, Nigeria
Upper Guinea: Ghana
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
spp
P.h.
All
322
6
46333
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
40 (5)
17
100
100
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:42 PM
Page 323
Ptychopetalum anceps
Oliv.
Olacaceae
Description
Guild: sb
Leaf: alternate, simple, oblong to elliptic, notophyll
(2.5-4 x 6.5-14 cm), herbaceous; subsessile
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

Ghana
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
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

Cte dIvoire, 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)
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
P.a.
119
33 (91)
40
52
24
23
77
18
45
45
All
46333
37
39
37
29
24
10
69
25
36
13
39
323
11/11/03
3:42 PM
Page 324
Puelia olyriformis
(Franch.) Clayton
Gramineae
Inflorescence: terminal and drooping, branched
(panicle, up to 15 cm long)
Flower: pale green
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)
Liberia (herbarium)
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
Leaf: simple, linear, microphyll (2-7 x 15-35 cm),
entire, herbaceous, smooth, glossy dark green
Data sources
FWTA, Van der Zon (1992)
spp
324
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
P.o.
67 (3)
20
60
20
20
60
100
60
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:42 PM
Page 325
Pycnanthus dinklagei
Warb.
Myristicaceae
Description
Guild: np
Max height: 25 m long (herbarium)
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
Distribution
Ghana
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).
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
P.d.
39
29 (28)
64
13
33
36
21
77
21
23
64
All
46333
37
39
37
29
24
10
69
25
36
13
39
325
11/11/03
3:43 PM
Page 326
Pyrenacantha glabrescens
(Engl.) Engl.
Icacinaceae
Description
Phenology
Guild: np
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
forest
Habitat
Most commonly found in established secondary
forest, and along forest borders (Kuzee pers.
comm., herbarium).
spp
326
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
65
35
29
71
69
25
36
13
39
P.g.
17
21 (14)
53
65
35
All
46333
37
39
37
29
24
10
Soil WHC
11/11/03
3:43 PM
Page 327
Raphia palma-pinus
(Gaertn.) Hutch.
Palmae
Description
Phenology
Guild: u
Life form: palm tree
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)
Fruit: fleshy, elongate-ovoid (4 cm x 8.8 cm),
orange-brown, with scales
Dispersal: by birds
Distribution
Data sources

Upper Guinea: Senegal, Gambia, Guinea, Sierra
Leone, Cte dIvoire, Ghana
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.
spp
Open (n)
R.p.
16
67 (3)
All
46333
37
Altitude
River
Coast
>500m
<2km
<5km
VW
56
37
44
12
44
56
19
31
44
39
37
29
24
10
69
25
36
13
39
0
4
Soil CMK
Soil WHC
327
11/11/03
3:43 PM
Page 328
Renealmia battenbergiana
Cummins ex Baker
Zingiberaceae
Description
Phenology
Guild: sb
Leaf: simple, elliptic, mesophyll (5-5.5 x 20 cm),
entire, coriaceous, reddish beneath
Inflorescence: at the top of leafy shoots, branched
(paniculate)
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)

Habitat
Found in moist places of dense forests (e.g. by
streams). On sandy soils (herbarium).
Regeneration
It tends to form clumps.
spp
328
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
R.b.
0 (3)
25
50
75
25
25
75
H
75
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:45 PM
Page 329
Renealmia longifolia
K.Schum.
Zingiberaceae
Description
Guild: u
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
Distribution
Upper Guinea: Liberia, Cte dIvoire (FWTA)
Distribution type: local, present in 3 30 cells,
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
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
R.l.
100 (1)
33
33
33
33
100
100
H
0
All
46333
37
39
37
29
24
10
69
25
36
13
39
329
11/11/03
3:45 PM
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
Leaf: alternate, simple, lanceolate to ovate,
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
Forest type: rainforest, evergreen forest,
deciduous forest, swamp forest, secondary forest
Data sources
FWTA, Mangenot (1957), Letouzey (1961),
spp
330
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
R.b.
37
21 (9)
70
46
54
16
57
27
11
H
62
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:45 PM
Page 331
Rinorea aylmeri
Chipp
Violaceae
Description
Phenology
Guild: u
Max. height: 5 m
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)
Fruit: dry dehiscent, triangular to globose (2 cm in
diameter), dark brown, sparsely puberulous to
glabrous
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
Distribution
Upper Guinea: Guinea-Bissau, Sierra Leone,
Liberia, Cte dIvoire, Ghana
Habitat
(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).
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
R.a.
36
22 (9)
42
14
37
17
43
83
17
53
H
31
All
46333
37
39
37
29
24
10
69
25
36
13
39
331
11/11/03
3:45 PM
Page 332
Sabicea arachnoidea
Hutch. & Dalziel
Rubiaceae
Description
Data sources
Guild: u
Leaf: opposite, simple, obovate, notophyll (4-5 x
10-12 cm), whitish beneath, pubescent above;
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
Other: a rather slender, pubescent climber.
Distribution
in 1 30 cell, distribution range is 22.7 km
Forest type: data unavailable.
Habitat
Data unavailable.
Phenology
spp
332
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.a.
- (0)
33
100
100
67
H
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:45 PM
Page 333
Sabicea bracteolata
Wernham
Rubiaceae
Description
Data sources
Guild: u
Leaf: opposite, simple, elliptic, notophyll (2.5-5 x
4-10 cm), entire, pubescent on the nerves below;
Flower: medium-sized; densely pubescent outside
Fruit: fleshy, subglobose or ellipsoid (2 cm or less
in diameter), soft and sweet, red; several seeds
Other: it probably starts as a shrub. The stems are
densely pubescent when young.
Distribution
Upper Guinea: Guinea
in 1 30 cell, only known from 3 collections in
Guinea
Habitat
Data unavailable.
Phenology
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.b.
- (0)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
333
11/11/03
3:45 PM
Sabicea discolor
Page 334
Stapf
Rubiaceae
Description
Phenology
Guild: pi
Life form: small winding climber
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
Distribution
FWTA, Hawthorne & Jongkind

(2004)

Forest type: montane rainforest, wet evergreen
forest, moist evergreen forest, swamp forest,
Habitat
rainfall, reaching an optimum around 2700 mm/yr
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).
spp
334
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
S.d.
61
82 (34)
10
33
18
33
28
31
75
23
49
36
All
46333
37
39
37
29
24
10
69
25
36
13
39
Data sources
11/11/03
3:45 PM
Page 335
Sabicea harleyae
Hepper
Rubiaceae
Description
Phenology
Guild: pi
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
Other: a slender climber with hairy stems.
Data sources
Distribution
FWTA, Hepper (1958), Hawthorne & Jongkind

(2004)

dIvoire
evergreen forest, secondary forest
Habitat
Practically only reported in disturbed open places
(e.g. roadsides, logged and open forests)
(herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.h.
14
73 (11)
29
36
58
36
100
79
21
All
46333
37
39
37
29
24
10
69
25
36
13
39
335
11/11/03
3:45 PM
Page 336
Salacia columna
N.Hall
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
Guild: sb
Life form: medium-sized irregularly winding woody
climber
Max. height: 10 m long (Hall 1962)
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
Distribution
Ghana
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).
spp
336
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
S.c
23
42 (12)
39
31
51
91
78
13
All
46333
37
39
37
29
24
10
69
25
36
13
39
Uses
The fruits are edible (herbarium).
Data sources
Hall (1962), Hall & Swaine (1981)
11/11/03
3:45 PM
Page 337
Salacia howesii
Hutch. & M.B.Moss
Celastraceae
Description
Habitat
Guild: np
Leaf: opposite, simple, mesophyll (5 x 14 cm),
entire, coriaceous, densely pubescent underneath
Flower: small (0.6 cm in diameter); yellowish; dishshaped
Fruit: fleshy, globose (approx. 2.5 cm in diameter),
orange
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
Phenology
Data unavailable.
Data sources
FWTA, Hall (1962), Hall & Lock (1975),
Hall & Swaine (1981)
Distribution
Continent: Nigeria, Gabon
range is 1985 km
forest, moist semi-deciduous forest
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.h.
25 (4)
50
25
75
50
50
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
337
11/11/03
3:45 PM
Page 338
Sapium aubrevillei
Leandri
Euphorbiaceae
Distribution
Ghana
List species (Vulnerable). It is rare (Hawthorne
1995a).
Habitat
Few records found in both dense and open forests.
We did not find any significant relationship with
environmental variables.
Regeneration
type (De la Mensbruge 1966).
Description
Phenology
Guild: pi
Max. diameter: 120 cm (FWTA)
(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
yellow; 2 seeds
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.
Data sources
Savill & Fox (1967), Hawthorne (1990, 1995a),
IUCN Red List (2000)
spp
338
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.a.
11
0 (7)
18
64
45
36
64
36
18
45
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:50 PM
Page 339
Scadoxus multiflorus (Martyn) Raf. ssp. longitubus (C.H.Wright) Friis & Nordal
Amaryllidaceae
Description
Guild: np
Life form: perennial bulbous ground herb
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)
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).
Hall & Swaine (1981), De Koning (1983)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.m.
60
42 (31)
48
35
47
82
15
35
H
58
All
46333
37
39
37
29
24
10
69
25
36
13
39
339
11/11/03
3:50 PM
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
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
forest, secondary forest. It can be locally abundant
in wet evergreen forest and in swamps of semideciduous forests (Hawthorne & Jongkind 2004).
Data sources
FWTA, Jenk (1969), Hall & Swaine (1981),
Habitat
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
spp
340
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.a.
47
33 (30)
51
60
10
27
53
43
49
H
43
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:50 PM
Page 341
Schizocolea linderi
(Hutch. & Dalziel) Bremek.
Rubiaceae
Description
Phenology
Guild: sb
Life form: small shrub
Leaf: opposite, simple, elliptic, notophyll (2-5 x 615.5 cm), entire, coriaceous, hairy midrib above;
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
Other: the stems, petioles and midribs are hairy.
Distribution
Data sources

forest
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).
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
S.l.
26
12 (17)
65
31
62
96
85
15
All
46333
37
39
37
29
24
10
69
25
36
13
39
341
11/11/03
3:50 PM
Page 342
(A.Chev.) Aubrv.
Rubiaceae
Description
Phenology
Guild: sb
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
range is 1086 km, Red List species (Vulnerable)
forest, semi-deciduous forest
Habitat
Found in the understorey, especially in wet places in
the forest (Savill & Fox 1967).
Regeneration
It regenerates in shade.
spp
342
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.p.
13
0 (10)
46
38
30
15
15
62
31
38
H
54
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/21/03
2:05 PM
Page 343
Scytopetalum tieghemii
Hutch. & Dalziel
Scytopetalaceae
Description
Regeneration
Guild: sb
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
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.
fruiting period in July (Savill & Fox 1967)
Distribution
Ghana
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
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)
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
M
S.t.
75
40 (40)
51
50
26
19
71
23
41
49
All
46333
37
39
37
29
24
10
69
25
36
13
39
343
11/11/03
3:51 PM
Page 344
Sericanthe adamii
(N.Hall) Robbr.
Rubiaceae
Description
Data sources
Guild: u
Life form: large woody climber, climbing with
recurved spines
Max. height: 35 m long
6-11 cm), entire, herbaceous, slightly hairy on the
nerves; with interpetiolar stipules
Flower: small; 8-merous; pedicel pubescent
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
Habitat
It occurs around 500 m altitude (Robbrecht 1981a).
Phenology
spp
344
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.a.
0 (2)
50
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:51 PM
Page 345
Sherbournia calycina
(G.Don) Hua
Rubiaceae
Description
Guild: pi
Max. height: 50 m long, 10 m high (herbarium)
7-18 cm), entire, coriaceous, sometimes with red
spots underneath; petiole first pubescent, thereafter
glabrous; interpetiolar stipules
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
Phenology
Distribution
Deciduousness: probably deciduous

Timing: flowering period from October to May;
fruiting period from November to August (De Koning
1983)

dIvoire, Ghana, Togo
Forest type: wet evergreen forest, gallery forest,
Habitat
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

type (cf. de Koning 1983).

(2004)
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
S.c.
125
44 (79)
51
46
26
24
68
30
36
46
All
46333
37
39
37
29
24
10
69
25
36
13
39
345
11/11/03
3:51 PM
Page 346
Sorindeia collina
Keay
Anacardiaceae
Description
Phenology
Guild: sb
Max. height: 10 m (Keay 1956)
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
Other: it has aerial roots.
Data sources
Distribution
FWTA, Keay (1956), Savill & Fox (1967), Hawthorne

& Jongkind (2004)

Forest type: montane high forest (FWTA). It is
locally common (herbarium).
Habitat
An understorey tree in montane high forests
(herbarium).
spp
346
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.c.
0 (3)
50
33
33
66
33
67
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:51 PM
Page 347
Soyauxia grandifolia
Gilg & Stapf
Medusandraceae
Description
Guild: sb
Leaf: alternate, simple, elongate to oblong,
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

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 Cte
dIvoire, it has been recorded on lateritic, loamy
sands, sandy clay, and sandy soils (herbarium).
Data sources
FWTA, Aubrville (1959), Hall & Swaine (1981),
spp
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
S.g.
19
44 (18)
47
21
26
26
42
74
21
47
11
32
All
46333
37
39
37
29
24
10
69
25
36
13
39
347
11/11/03
3:51 PM
Page 348
Strephonema pseudocola
A.Chev.
Combretaceae
Description
(e.g. rivers, waterfalls, and even swamps). Often on

Guild: sb
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
Timing: flowering period from August to February;
fruiting period from October to March
Distribution
Ghana
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
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)
spp
348
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.p.
53
20 (25)
51
40
38
19
13
66
21
32
H
47
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:53 PM
Page 349
Strombosiopsis nana
Breteler
Olacaceae
Description
Phenology
Guild: u
Life form: shrub, treelet
Max. height: 3 m
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
Other: the branchlets are green and slightly
compressed to narrowly winged.
Data sources
Distribution
FWTA, Breteler (2001), Hawthorne

& Jongkind (2004)

in 2 30 cells. Recorded only in Liberia.
Habitat
Found in rainforest at altitudes between 0-100 m
(Breteler 2001).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.n.
0 (1)
33
67
33
100
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
349
11/11/03
3:53 PM
Page 350
Strychnos dinklagei
Gilg
Loganiaceae
Regeneration
It regenerates in primary forest (De Rouw 1991)
Phenology
Description
Distribution
Guild: np
3-13 cm), revolute, coriaceous, glabrous on both
sides
(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.

Upper Guinea: Guinea, Liberia, Cte dIvoire, Ghana
Forest type: rainforest, coastal forest, mangroves,
secondary forest
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).
spp
350
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
S.d.
50
46 (28)
10
58
14
60
10
36
44
20
18
34
28
All
46333
37
39
37
29
24
10
69
25
36
13
39
Data sources
FWTA, Leeuwenberg (1969), De Rouw (1991)
11/11/03
3:53 PM
Page 351
Strychnos melastomatoides
Gilg
Loganiaceae
Description
Guild: u
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
Distribution type: continuous, local, present
Forest type: rainforest, gallery forest
Phenology
Habitat
It occurs along riverbanks or in gallery forest,
at low elevations (Leeuwenberg 1969).
Data sources
FWTA, Leeuwenberg (1969), Hawthorne & Jongkind
(2004)
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.m.
- (0)
43
57
43
57
43
14
14
H
43
All
46333
37
39
37
29
24
10
69
25
36
13
39
351
11/11/03
3:53 PM
Page 352
Strychnos millepunctata
Leeuwenb.
Loganiaceae
Description
Distribution
Guild: sb
Life form: medium-sized woody climber
Leaf: opposite, simple, obovate, notophyll (2-8 x
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.

species (Vulnerable)
Forest type: rainforest, coastal forest, riverine
Habitat
It is usually found on riverbanks, on sandy soils
(herbarium).
Regeneration
It regenerates in primary forest (De Rouw 1991).
Phenology
Data sources
Leeuwenberg (1969), De Rouw (1991), IUCN Red
List (2000)
spp
352
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.m.
14
18 (11)
14
36
36
64
36
57
43
36
21
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:53 PM
Page 353
Strychnos odorata
A.Chev.
Loganiaceae
Description
Phenology
Guild: u
Max. height: 40 m high, 100 m long (herbarium)
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.
Data sources
Distribution
FWTA, Leeuwenberg (1969)

Habitat
Found in moist places in rainforest at low elevations,
usually, on riverbanks (Leeuwenberg 1969).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.o.
0 (2)
75
100
50
50
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
353
11/11/03
3:54 PM
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
Regeneration
type (cf. De Koning 1983), and regenerates in the
shade (Hall & Swaine 1981).
Description
Phenology
Guild: np
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.

Timing: flowering period from December to April;
fruiting period in December (De Koning 1983)
Data sources

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)
Leeuwenberg (1969), Hall & Swaine

(1981), De Koning (1983)
spp
354
Open (n)
S.s.
101
53 (70)
All
46333
37
Altitude
River
Coast
>500m
<2km
<5km
VW
46
11
44
35
11
69
24
31
58
39
37
29
24
10
69
25
13
39
9
4
Soil CMK
Soil WHC
36
11/11/03
3:54 PM
Page 355
Suregada ivorensis
(Aubrv. & Pellegr.) J.Lonard
Euphorbiaceae
Description
Phenology
Guild: u
Leaf: alternate, simple, entire
Inflorescence: data unavailable
Distribution
Upper Guinea: southeast Cte dIvoire
in 2 30 cells. The species is endangered (C.
Chatelain, pers. comm.).
forest
Data sources
Lonard (1958), Aubrville (1959),
Habitat
Data unavailable.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
S.i.
100 (1)
100
50
50
50
50
50
H
50
All
46333
37
39
37
29
24
10
69
25
36
13
39
355
11/11/03
3:54 PM
Page 356
Tabernaemontana africana
Hook.
Apocynaceae
Forest type: wet evergreen forest, moist semideciduous forest, dry semi-deciduous forest
Habitat
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
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),
Distribution
spp
356
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
T.a.
171
36 (33)
36
11
17
33
21
29
60
37
29
11
39
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:54 PM
Page 357
Talbotiella gentii
Hutch. & Greenway
Leguminosae-Caes.
Description
Habitat
Guild: sb
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
Distribution
Upper Guinea: Ghana
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
Ghanas 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
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.g.
23
10 (20)
13
30
74
26
22
78
22
48
All
46333
37
39
37
29
24
10
69
25
36
13
39
357
11/11/03
3:54 PM
Page 358
Tapinanthus belvisii
(DC.) Danser
Loranthaceae
Description
Phenology
Guild: pi
Life history: hemi-parasitic shrub
Max. height: 1 m
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)
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 & Hall (1961), Polhill & Wiens (1998)
Distribution
1998)
(herbarium, Polhill & Wiens 1998)
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).
spp
358
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.b.
20
50 (2)
25
65
15
60
25
45
35
20
45
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:56 PM
Page 359
Tapinanthus praetexta
Polhill & Wiens
Loranthaceae
Description
Phenology
Guild: pi
Max. height: 0.7 m (Polhill & Wiens 1998)
Leaf: opposite, simple, elliptic to lanceolate,
microphyll (2-5 x 5-9 cm), entire, herbaceous
Inflorescence: axillary, umbel (with 8-12 flowers)
corolla tube pale pink, lobes red to purple
red; 1 seed
Seed: medium-sized
Dispersal: by birds
Timing: flowering period from July to February
Distribution
1998)
Upper Guinea: Cte dIvoire (Polhill & Wiens 1998)
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 Cte
dIvoire (Polhill & Wiens 1998).
Forest type: coastal forest
Data sources
Polhill & Wiens (1998)
Habitat
Data unavailable.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.p.
11
100 (2)
91
45
55
64
18
18
55
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
359
11/11/03
3:56 PM
Tapura fischeri
Page 360
Engl.
Dichapetalaceae
Description
Regeneration
Guild: u
Max. height: 20 m (Breteler 1986)
Max. diameter: 45 cm (Breteler 1986)
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)
Breteler 1986)
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).
spp
360
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.f.
11
0 (6)
90
55
45
18
45
27
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:56 PM
Page 361
Tapura ivorensis
Breteler
Dichapetalaceae
Description
Phenology
Guild: sb
Max. height: 10 m
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.
Data sources
Breteler (1986), Hawthorne (1990, 1995a), IUCN
Red List (2000), Hawthorne & Jongkind (2004)
Distribution
in 2 30 cells, Red List species (Vulnerable)
Habitat
It occurs especially along streambanks
(Hawthorne 1995a).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.i.
0 (1)
33
67
33
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
361
11/11/03
3:56 PM
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.

(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
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;
Inflorescence: terminal, laxly branched cyme
Flower: small (corolla tube 1 cm long, lobes 0.7
cm); tube-shaped, 5 lobes, white
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
type (cf. De Koning 1983).
Phenology
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, Cte
dIvoire and Ghana. It has its largest population in
Upper Guinea and is present in 27 30 cells. It
spp
362
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.v.
66
22 (36)
61
36
53
10
80
20
22
78
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:56 PM
Page 363
Tetraberlinia tubmaniana
J.Lonard
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
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
(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
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

Upper Guinea: Liberia (Wieringa 1999)
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).
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),
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
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.t.
25
11 (9)
92
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
363
11/11/03
3:56 PM
Page 364
Thomandersia anachoreta
Heine
Acanthaceae
Description
Phenology
Guild: np
Life form: treelet or shrub
Max. height: 2 m (Heine 1966)
Leaf: opposite, simple, lanceolate, notophyll (3-5 x
(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

Upper Guinea: Liberia, Cte dIvoire (Heine 1966,
herbarium)
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).
spp
364
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.a.
33 (6)
14
43
43
58
14
86
29
14
57
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:56 PM
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
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
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
d'Ivoire, Ghana.
Distribution type: continuous distribution,
widespread, distribution range is > 1411 km
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
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
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.l.
71
39 (44)
35
18
13
50
24
13
76
17
39
11
42
All
46333
37
39
37
29
24
10
69
25
36
13
39
365
11/11/03
3:56 PM
Page 366
Trichilia djalonis
A.Chev.
Meliaceae
Description
Phenology
Guild: u
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

Upper Guinea: Guinea, Liberia, Cte dIvoire
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).
spp
366
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.d.
12
33 (9)
67
50
33
50
67
33
42
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:56 PM
Page 367
Trichilia megalantha
Harms
Meliaceae
Description
Phenology
Guild: np
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.

Jongkind 2004)
Data sources
FWTA, De Wilde (1968), Hall & Swaine (1981),
Distribution
Continent: Guinea wide: from Cte dIvoire to
Nigeria (Hall & Swaine 1981), Benin, Nigeria
(herbarium)
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.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.m.
17 (6)
50
63
13
25
75
25
25
75
All
46333
37
39
37
29
24
10
69
25
36
13
39
367
11/11/03
3:56 PM
Page 368
Trichilia ornithothera
J.J.de Wilde
Meliaceae
Habitat
Species occurrence increases significantly in places
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
Jongkind 2004)
Timing: flowering period from June to September;
fruiting period from December to April (De Koning
1983)
Description
Guild: np
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

dIvoire, Ghana
forest
The fruits are boiled and used in local medical

treatments (herbarium).
Data sources
FWTA, De Wilde (1968), De Koning (1983),
spp
368
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.o.
30
67 (21)
47
13
40
27
16
90
40
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:58 PM
Page 369
Trichoscypha laxissima
Breteler
Anacardiaceae
Description
Guild: u
Life form: small shrub
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
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)
Environmental table unavailable.

There were no sufficient data on the environmental conditions of this single specimen.
369
11/11/03
3:58 PM
Page 370
Triclisia dictyophylla
Diels
Menispermaceae
Description
Habitat
Guild: pi
Max. height: 7 m high, 20 m long (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
Cte dIvoire to the Democratic Republic of Congo
(herbarium)
Distribution type: continental disjunct, widespread,
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).
spp
370
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.d.
12
33 (6)
17
75
16
42
58
33
58
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:58 PM
Page 371
Tristemma akeassii
Jacq.-Fl.
Melastomataceae
Description
Phenology
Guild: np
Life form: erect or creeping herb, rather woody at
the base
Max. height: 1.5 m (De Koning 1983)
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-Flix (1976), De Koning (1983)
Distribution
Continent: Nigeria (Jacques-Flix 1976)
Forest type: montane forest, evergreen forest,
secondary forest. It is locally abundant (herbarium).
Habitat
Species occurrence increases significantly in places
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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.a
39
57 (28)
44
11
46
25
19
56
38
31
51
All
46333
37
39
37
29
24
10
69
25
36
13
39
371
11/11/03
3:58 PM
Page 372
Tristemonanthus nigrisilvae
(N.Hall) N.Hall
Celastraceae
Description
Phenology
Guild: sb
5-10 cm), often serrate towards the apex,
coriaceous
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

Forest type: wet evergreen forest, coastal forest,
secondary forest (herbarium)
Hall (1958, 1959), De Koning (1983), De Rouw

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).
spp
372
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.n.
15
21 (14)
27
47
47
87
13
60
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:58 PM
Page 373
Turraea ghanensis
J.B.Hall
Meliaceae
Description
Phenology
Guild: sb
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

Jongkind 2004)
Distribution
Upper Guinea: Cte dIvoire, Ghana. Described as
a Ghanaian endemic (Hall 1976). There is one
doubtful record from Cte dIvoire.
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.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.g.
0 (6)
14
100
29
71
29
All
46333
37
39
37
29
24
10
69
25
36
13
39
373
11/11/03
3:59 PM
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

Jongkind 2004)
Timing: flowering not clearly seasonal (Hall &
Swaine 1981)
Guild: sb
Life form: pigmy tree or shrub
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
Cte dIvoire, it has been recorded in rainforest,
gallery forest, savanna (forest and cultivated), and
secondary forests (herbarium).
Continent: from Sierra Leone to Benin

Forest type: it occurs in a
Data sources
FWTA, Hall & Swaine (1981), Hawthorne & Jongkind
(2004)
spp
374
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
T.h.
52
29 (21)
31
12
67
16
13
13
54
33
31
23
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:59 PM
Page 375
Uapaca chevalieri
Beille
Euphorbiaceae
Description
Phenology
Guild: sb
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.
Distribution
Uses

dIvoire (herbarium, FWTA)
Forest type: moist evergreen forest, swamp forest,
gallery forest
It has edible fruits.
Data sources
FWTA, Thoen & Ducousso (1989),
Habitat
Found in swamps and riversides of montane regions
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
U.c.
12
18 (11)
75
58
75
25
25
75
17
H
0
All
46333
37
39
37
29
24
10
69
25
36
13
39
375
11/11/03
3:59 PM
Page 376
Uapaca paludosa
Aubrv. & Landri
Euphorbiaceae
Habitat
In swamps in evergreen forests, in lowland regions
Regeneration
type (cf. De la Mensbruge 1966).
Phenology
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
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
Other: it has stilt roots. Similar, particularly as a
sapling, to U. guineensis. The wood density is 0.76
g/cm3.
Uses
Distribution

Continent: Nigeria (FWTA), Cameroon, Gabon
(herbarium)
Upper Guinea: Liberia, Cte dIvoire (herbarium),
Ghana
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)
spp
376
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
U.p.
0 (4)
11
56
11
22
67
33
67
11
11
44
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
3:59 PM
Page 377
Uvaria dinklagei
Engl. & Diels
Annonaceae
Description
Phenology
Guild: u
Life form: irregularly winding woody climber
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
Other: the twigs are hairy.
Distribution
Uses

in 1 30 cell
Forest type: coastal shrubland
Data sources
Habitat
Data unavailable.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
U.d.
100 (1)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
377
11/11/03
3:59 PM
Uvaria ovata
Page 378
(Dunal) DC.
Annonaceae
Phenology
Distribution
Ghana, Togo
Distribution type: continuous, widespread,
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 Cte dIvoire, found in savanna and
secondary regrowth (herbarium).
Description
Guild: np
Life form: irregularly winding woody climber
Leaf: alternate, simple, ovate to elliptic to
lanceolate, microphyll (1.5-5 x 4-11 cm), entire,
coriaceous, pubescent below, aromatic
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
Other: the twigs are pubescent.
Data sources
FWTA, Hall & Swaine (1981), Ak Assi (1984),
Habitat
It forms dense thickets in dry areas along
roadsides, near the coast, and grasslands.
Regeneration
spp
378
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
U.o.
34
78 (9)
98
82
18
56
H
24
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
4:02 PM
Page 379
Uvaria sassandrensis
Jongkind
Annonaceae
Description
Phenology
Guild: u
Leaf: alternate, simple, entire, coriaceous
Flower: large; calyx pale green; corolla pinkish to
orange
Fruit: yellow
Other: a new species (Jongkind in prep.). Starts as
a shrub when young. The branchlets and petioles
are red-brown pilose.
Data sources
Distribution

Forest type: secondary forest (herbarium)
Habitat
Usually found in the understorey or in secondary
forests. Sometimes on riverbanks. On sandy or
loamy soils (herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
U.s.
10
44 (9)
40
30
80
20
90
10
60
H
40
All
46333
37
39
37
29
24
10
69
25
36
13
39
379
11/11/03
4:02 PM
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
Habitat
It is found in the forest understorey (herbarium).
Regeneration
Phenology
Description
Data sources
Guild: sb
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.
Distribution
Upper Guinea: Cte dIvoire, Ghana. The record
from Cte dIvoire is doubtful.
spp
U.g.
All
380
4
46333
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
0 (4)
25
75
25
75
25
50
37
39
37
29
24
10
69
25
36
13
39
11/11/03
4:02 PM
Page 381
Vahadenia caillei
A.Chev.
Apocynaceae
Description
Phenology
Guild: np
Max. height: 40 m long (Haegens 1994)
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
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),

Data sources

dIvoire (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.
Forest type: high forest, fringe or secondary
forest, swamp forest
Habitat
(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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
V.c.
25
60 (5)
44
32
28
44
28
44
48
36
H
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
381
11/11/03
4:02 PM
Page 382
Vangueriella vanguerioides
(Hiern) Verdc.
Rubiaceae
Description
Phenology
Guild: np
Max. height: 10 m high (herbarium)
Leaf: opposite, simple, lanceolate to obovate,
notophyll (2-5 x 4.5-11.5 cm), entire, herbaceous,
pubescent when young; interpetiolar stipules
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
dIvoire, Ghana (herbarium, Verdcourt 1987)
dry forest, savanna, gallery forest, secondary
forest, disturbed forest
Habitat
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).
spp
382
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
L
V.v.
41
33 (21)
29
44
37
19
40
68
32
39
37
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
4:02 PM
Page 383
Vepris tabouensis
(Aubrv. & Pellegr.) W.Mziray
Rutaceae
Description
Phenology
Guild: u
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.

January; fruiting period from September to October
(Voorhoeve 1965)
Distribution
Data sources
Continent: Nigeria
forest
Voorhoeve (1965),
Mziray (1995)
Habitat
Sometimes found close to streams or rivers
(herbarium).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
V.t.
12 0 (7)
42
33
25
41
100
67
33
All
46333
37
39
37
29
24
10
69
25
36
13
39
383
11/21/03
5:24 PM
Page 384
Vernonia titanophylla
Brenan
Compositae
Distribution
Continent: Cameroon, Equatorial Guinea, Gabon,
Democratic Republic of Congo (herbarium)
Upper Guinea: Guinea Bissau (Moutsambout
1990), Liberia, Cte dIvoire, Ghana (herbarium)
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
Max. height: 10 m (Moutsambout 1990)
Max. diameter: 30 cm (Moutsambout 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

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.
Moutsambout (1990), Hawthorne (1995a),

spp
384
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
V.t.
75 (4)
60
40
60
40
60
40
40
20
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
4:02 PM
Page 385
Whitfieldia colorata
C.B.Clarke ex Stapf
Acanthaceae
Description
Phenology
Guild: np
Life form: shrub, sometimes scrambling
Leaf: opposite, simple, lanceolate, mesophyll
(4.5-11 x 12-27 cm), entire, coriaceous, glabrous
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
Other: it has dark green stems, with some purple
above the nodes. It has prominent purple midribs.
Data sources
FWTA
Distribution
forest, savanna, secondary forest
Habitat
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).
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
W.c.
40
41 (17)
60
15
20
10
62
80
18
60
23
All
46333
37
39
37
29
24
10
69
25
36
13
39
385
11/11/03
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)
Ghana (herbarium)
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
Habitat
Sometimes found along streams (herbarium).
Data sources
Description
Regeneration
Guild: np
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
Open (n)
Altitude
River
Coast
Soil CMK
>500m
<2km
<5km
VW
Soil WHC
M
X.e.
24
8 (12)
13
25
33
46
16
50
46
42
29
All
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
4:02 PM
Page 387
Xylopia elliotii
Engl.
Annonaceae
Description
Phenology
Guild: sb
Max. height: 9 m (FWTA, herbarium)
Leaf: alternate, simple, lanceolate, microphyll (2-3 x
5-9 cm), entire, pubescent beneath
Flower: medium-sized (approx. 2.5 cm long); white
to yellowish; sweet scented
Other: it has hairy twigs.
Data sources
FWTA, Savill & Fox (1967), Hawthorne (1995a),
Distribution
dIvoire
species (Vulnerable)
(herbarium). In Sierra Leone, it has been recorded in
fringing forest (Savill & Fox 1967).
Habitat
It occurs in the forest understorey.
X
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
X.e.
0 (2)
11
44
11
44
44
56
44
44
11
All
46333
37
39
37
29
24
10
69
25
36
13
39
387
11/11/03
4:02 PM
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
Regeneration has been found to be obviously
depleted in burnt forest (Hawthorne 1994).
Phenology
Timing: flowering period from November to April
(De Koning 1983); fruiting period all year (De Koning
1983)
Description
Guild: sb
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
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
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
28
14 (14)
61
11
43
43
71
25
25
71
46333
37
39
37
29
24
10
69
25
36
13
39
11/11/03
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Page 389
(Ak Assi) Waterman
Rutaceae
Description
Phenology
Guild: pi
Max. height: 15 m (Ak Assi 1960)
Max. diameter: 2 cm (Ak 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
Ak Assi (1960), IUCN Red List (2000), Hawthorne
& Jongkind (2004)
Distribution
Upper Guinea: Cte dIvoire (herbarium, Ak Assi
1960)
species (Endangered)
Forest type: coastal forest (Ak Assi 1960)
Habitat
Data unavailable.
spp
Open (n)
Altitude
River
Coast
>500m
<2km
<5km
VW
Soil CMK
L
Soil WHC
L
Z.p.
100 (1)
100
100
100
All
46333
37
39
37
29
24
10
69
25
36
13
39
Z
389
390
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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
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 Cte dIvoire
(Chatelain et al., chapter 2), and a little bit later in the
neighbouring countries. Especially in Sierra Leone and
Cte dIvoire 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
(Aubrville 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 Fort Classe Haut Sassandra in Cte dIvoire.
Species data
Species selection
The focus of this chapter is on the large commercial
tree species of Sierra Leone, Liberia, Cte dIvoire 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
Cte dIvoire 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 Cte dIvoire
we drew largely on data collected by SODEFOR (Clment
& Guinaudeau 1973, SODEFOR 1978, 1979) and for
Ghana the national forest inventory data were used (Wong
1989, Hawthorne 1995a). In Liberia and Cte dIvoire
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 Cte dIvoire 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 Cte
dIvoire. 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
(Aubrville 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,
11/11/03
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Page 393
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
11/11/03
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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
11/11/03
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Page 395
Albizia ferruginea
(Guill. & Perr.) Benth.
Leguminosae-Mim.
A
Description
Guild: np
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
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
(1981), Kyereh et al. (1993), Kyereh (1994),
Table data are not available.
395
11/11/03
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Alstonia boonei
Page 396
De Wild.
Apocynaceae
A
Description
Regeneration
Guild: pi
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)
Forest type: moist evergreen forest, upland
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
A.b.
151
27
58
29
61
10
21
72
All sites
198
26
57
31
58
11
25
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
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
Harms
Leguminosae-Pap.
A
Description
Regeneration
Guild: np
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)
November; fruiting period from January to February
(Voorhoeve 1965)
Distribution
Continent: Benin to Cameroon, Congo (Brazzaville),
Sudan
Upper Guinea: Guinea to Togo
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)

(1981), Hawthorne (1995a), Swaine (1996)
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
A.p.
153
24
59
31
59
10
23
70
All sites
198
26
57
31
58
11
25
72
397
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Page 398
Anopyxis klaineana
(Pierre) Engl.
Rhizophoraceae
A
Description
Regeneration
Guild: np
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.

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
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
(1981), Hawthorne (1995a), IUCN Red List (2000)
Spp
398
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
A.k.
103
15
59
13
14
42
49
10
35
57
All sites
198
26
57
31
58
11
25
72
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Page 399
Anthonotha fragrans
(Baker f.) Exell & Hillcoat
Leguminosae-Caes.
A
Description
Regeneration
Guild: np
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.

type (cf. Voorhoeve 1965).
Phenology
Dispersal: no obvious means of dispersal
(Hawthorne 1995a)
Timing: flowering period in December; fruiting
period in April (Taylor 1960)
Distribution
1965)
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
(1981), Alexander (1989), Hawthorne (1995a)
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
A.f.
63
11
11
49
19
21
63
30
57
37
All sites
198
26
57
31
58
11
25
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
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
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)
Spp
400
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
>400m
VW
Soil CMK
L
Soil WHC
L
A.t.
159
27
60
27
60
13
20
71
All sites
198
26
57
31
58
11
25
72
Data sources
Taylor (1960), Osmaston (1965), Voorhoeve (196),
Berg (1977, 1978), Hall & Okali (1979), Hall &
Swaine (1981), Hawthorne (1995a), Swaine (1996)
11/11/03
4:07 PM
Page 401
Berlinia confusa
Holy
Leguminosae-Caes.
Description
Guild: sb
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.
Distribution
Continent: Benin to Gabon (Voorhoeve 1965)
1965)
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

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).

(1981), De Koning (1983), Hawthorne (1995a)
Phenology
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
B.spp.
140
21
62
36
56
25
H
68
All sites
198
26
57
31
58
11
25
72
401
11/11/03
4:07 PM
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
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
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)
Spp
402
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
C.s.
128
17
61
12
10
38
52
30
63
All sites
198
26
57
31
58
11
25
72

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.)
11/11/03
4:07 PM
Page 403
Ceiba pentandra
(L.) Gaertn.
Bombacaceae
Description
Habitat
Guild: pi
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
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)
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.).
semi-deciduous forest, secondary forest (Hall &
Swaine 1981)
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).

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
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
C.p.
167
27
58
30
58
12
22
69
All sites
198
26
57
31
58
11
25
72
403
11/11/03
4:11 PM
Page 404
Celtis adolfi-friderici
Engl.
Ulmaceae
Description
Guild: pi
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
Other: a slender tree with narrow, high buttresses
and a rounded, dark crown. The boughs are
horizontal and drooping at the tips.
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: Cte dIvoire, 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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
C.spp.
154
29
58
28
60
12
23
69
All sites
198
26
57
31
58
11
25
72
Data sources
Hall & Swaine (1981), Hawthorne (1995a)
11/11/03
4:11 PM
Page 405
Celtis mildbraedii
Engl.
Ulmaceae
Description
Regeneration
Guild: sb
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
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: Cte dIvoire, Ghana, Togo (Hall &
Swaine 1981)
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).
Growth
In Tropical Shelterwood System plots, height
increments of 1-3 m, in 4 years, have been
observed (Taylor 1960).
Phenology
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
C.spp.
154
29
58
28
60
12
23
69
All sites
198
26
57
31
58
11
25
72
405
11/11/03
4:11 PM
Page 406
Chrysophyllum giganteum
A.Chev.
Sapotaceae
Description
Phenology
Guild: sb
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)
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.
Distribution
Uses
Continent: Benin to Nigeria (Hall & Swaine 1981)

Upper Guinea: Sierra Leone to Togo (Hall &
Swaine 1981)
semi-deciduous forest (Savill & Fox 1967, Hall &
Swaine 1981). It is common in Ghana (Hall &
Swaine 1981).
Regeneration
type (cf. De la Mensbruge 1966).
406
Data sources
Savill & Fox (1967), Hall & Swaine (1981),
Hawthorne (1990)
11/11/03
4:12 PM
Page 407
Daniellia ogea
(Harms) Rolfe ex Holland
Leguminosae-Caes.
Description
Regeneration
Guild: pi
leaflets, obovate to oblong, microphyll (1-3.5 x 3-9
cm), entire; leaves and leaflets of saplings and
water shoots much larger
(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.

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).
Phenology
January; fruiting period from January to March
(Voorhoeve 1965)
Distribution
forest (Voorhoeve 1965, Hall & Swaine 1981). It is
common in Ghana (Hall & Swaine 1981).
Data sources
Habitat

(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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
D.spp
134
21
58
11
10
37
55
27
67
All sites
198
26
57
31
58
11
25
72
407
11/11/03
4:12 PM
Page 408
Daniellia thurifera
Benn.
Leguminosae-Caes.
Description
Regeneration
Guild: pi
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.

type (De Koning 1983). It regenerates in shade and
small gaps (Hall & Swaine 1981).
Phenology
Deciduousness: deciduous in October to
November (Savill & Fox 1967)
December (De Koning 1983); fruiting period from
March to April (Savill & Fox 1967
Distribution
1981)
Leone, Liberia, Cte dIvoire, Ghana (Hall & Swaine
1981)
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
Uses
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
D.spp
134
21
58
11
10
37
55
27
67
All sires
198
26
57
31
58
11
25
72
Data sources
FWTA, Taylor (1960), Savill & Fox (1967),
Voorhoeve (1965), Hall & Swaine (1981), De Koning
(1983)
11/11/03
4:12 PM
Page 409
Distemonanthus benthamianus
Baill.
Leguminosae-Caes.
Description
Guild: np
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.
Distribution
1965)
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

Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
D.b
139
25
61
32
58
10
23
71
All sites
198
26
57
31
58
11
25
72
409
11/11/03
4:12 PM
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
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. angolense as
the most shade-bearing of the Entandrophragmas.
Common, particularly in better-drained sites.
Description
Regeneration
Guild: np
Max. height: > 50 m (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
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)
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 Cte dIvoire in a 3 ha subsample of a
plot containing several adults (De Klerk 1991).
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
E.spp.
159
28
61
28
62
10
20
74
All sites
198
26
57
31
58
11
25
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)
11/11/03
4:12 PM
Page 411
Entandrophragma candollei
Harms
Meliaceae
Description
Regeneration
Guild: np
Leaf: paripinnately compound, 10-20 opposite
leaflets, elliptic, oblong or obovate, notophyll (2.5-6
x 5-16 cm), entire to undulate
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 Cte dIvoire in a 3 ha subsample of a plot
containing several adults (De Klerk 1991).
Distribution
evergreen forest, moist semi-deciduous forest (Hall
& Swaine 1981). It is common in Ghana (Hall &
Swaine 1981). A Red List species (Vulnerable).
Habitat
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
Growth rings in this species are likely to be annual

Phenology
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
E.spp.
159
28
61
28
62
10
20
74
All sites
198
26
57
31
58
11
25
72
411
11/11/03
4:12 PM
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.
Distribution
Continent: Benin to Uganda, Congo (Voorhoeve
1965)
semi-deciduous forest. A Red List species
(Vulnerable).
Habitat
(regression analysis).
Regeneration
Description
Spp
412
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
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

(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
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 Cte dIvoire in a 3 ha
subsample of a plot containing several adults (De
Klerk 1991).
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
E.spp.
159
28
61
28
62
10
20
74
All sites
198
26
57
31
58
11
25
72
Pieters (1958), Sawyerr (1960), Taylor (1960), Keay

(1961), Voorhoeve (1965), De la Mensbruge (1966),
De Klerk (1991), Hawthorne (1995a), IUCN Red List
(2000)
11/11/03
4:12 PM
Page 413
Entandrophragma utile
(Dawe & Sprague ) Sprague
Meliaceae
Description
Guild: np
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
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.
Distribution
Continent: Benin to Uganda, Angola (Voorhoeve 1965)
Forest type: upland evergreen forest, moist semideciduous forest, dry semi-deciduous forest. A Red
List species (Vulnerable).
Habitat
(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
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
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
E.spp.
159
28
61
28
62
10
20
74
All sites
198
26
57
31
58
11
25
72
413
11/11/03
4:13 PM
Page 414
Erythrophleum ivorense
A.Chev.
Leguminosae-Caes.
Description
Regeneration
Guild: np
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

type (cv. Voorhoeve 1965). There is little
regeneration (Taylor 1960).
Phenology
Timing: flowering period from May to September
(Voorhoeve 1965)
Distribution
Ghana, Togo (Voorhoeve 1965)
forest, moist semi-deciduous forest, high forest,
secondary forest
The abundance of Erythrophleum spp. as a group

declines with soil fertility (regression analysis).
414
The wood is very hard and is used for railway

sleepers and bridging timber (Voorhoeve 1965).
Data sources
Habitat
Spp
Uses
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
E.spp.
121
27
49
12
12
42
46
12
34
57
All sites
198
26
57
31
58
11
25
72

(1981), Lock (1989), Hawthorne (1995a)
11/11/03
4:13 PM
Page 415
Erythrophleum suaveolens
(Guill. & Perr.) Brenan
Leguminosae-Caes.
Description
Regeneration
Guild: np
Max. height: > 25 m (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)
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)
Distribution
Continent: widespread in Africa, Asia (Lock 1989)
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
E.spp.
121
27
49
12
12
42
46
12
34
57
All sites
198
26
57
31
58
11
25
72
415
11/11/03
4:13 PM
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
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, 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
Max. height: 30 m (Zwetsloot 1981)
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).
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
Regeneration
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
F.a.
142
25
61
31
58
11
22
71
All sites
198
26
57
31
58
11
25
72
11/11/03
4:13 PM
Page 417
Gilbertiodendron preussii
(Harms) J.Lonard
Leguminosae-Caes.
Description
Regeneration
Guild: sb
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
(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.

(cf. Voorhoeve 1965).
Phenology
Dispersal: seeds are ejected from the pod
(Voorhoeve 1965)
December; fruiting period from July to September
(Voorhoeve 1965)
Distribution
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
G.p.
48
54
21
21
63
31
65
31
All sites
198
26
57
31
58
11
25
72
417
11/11/03
4:13 PM
Guarea cedrata
Page 418
(A.Chev.) Pellegr.
Meliaceae
Distribution
Continent: Benin to Uganda, Congo (Voorhoeve
1965)
semi-deciduous forest, dry semi-deciduous forest
(Hall & Swaine 1981). A Red List species
(Vulnerable).
Habitat
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
Max. diameter: 100 cm above the buttresses
(Voorhoeve 1965)
(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
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
>400m
VW
Soil CMK
L
Soil WHC
L
G.c.
136
22
67
10
29
61
10
19
74
All sites
198
26
57
31
58
11
25
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
Phenology
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),
Bada (1989), Kyereh et al. (1993), Hawthorne
(1995a), Swaine (1996), Swaine et al. (1997), IUCN
Red List (2000)
11/11/03
4:13 PM
Page 419
Guibourtia ehie
(A.Chev.) J.Lonard
Leguminosae-Caes.
Description
Guild: np
Leaf: alternate, pinnately compound, 2 leaflets,
obliquely elliptic, microphyll (1.5-3.5 x 3-8 cm),
glossy green above, paler beneath
(spike, 4-9 cm long)
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.
Distribution
Continent: Upper Guinea endemic (Voorhoeve
1965)
(Voorhoeve 1965)
semi-deciduous forest (Hall & Swaine 1981). It is
common in Ghana (Hall & Swaine 1981). A Red List
Habitat
Its abundance increases with altitude and decreases
with rainfall (regression analysis). The species is
shade tolerant (Taylor 1960).
Phenology
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
(1966), Hall & Swaine (1981), Hawthorne (1995),
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
G.e.
88
33
59
35
58
23
74
All sites
198
26
57
31
58
11
25
72
419
11/11/03
4:13 PM
Page 420
Hallea ledermannii
(K.Krause) Verdc.
Rubiaceae
Description
Habitat
Guild: sw
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

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
Dispersal: by wind
January; fruiting period from February to May
(Voorhoeve 1965)
Data sources
Taylor (1960), Voorhoeve (1965), Jenk (1969),
Osain (1973), IUCN Red List (2000)
Map data are not available.
420
11/11/03
4:13 PM
Page 421
Khaya anthotheca
(Welw.) C.DC.
Meliaceae
Description
Regeneration
Guild: np
leaflets, leaflet elliptic, oblong or obovate (2.5-5.5 x
Inflorescence: terminal, branched (raceme or
panicle, 6-25 cm long)
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
Voorhoeve 1965). Very sparse regeneration of this
species was found in Cte dIvoire in a 3 ha
subsample of a plot containing several adults (De
Klerk 1991).
Distribution
Continent: Benin to Uganda, Angola (Voorhoeve
1965)
dIvoire, Ghana, Togo (Voorhoeve 1965)
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
Phenology
Deciduousness: evergreen or briefly
deciduous (Voorhoeve 1965)
Timing: flowering period from September
to October and January to February; fruiting
period in January (Voorhoeve 1965)
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
K.spp.
151
27
64
27
62
11
19
75
All sites
198
26
57
31
58
11
25
72
421
11/11/03
4:13 PM
Page 422
Khaya grandifoliola
C.DC.
Meliaceae
Habitat
analysis).
Regeneration
Guild: np
opposite or sub-opposite leaflets, elliptic to oblongelliptic (7 x 16 cm), entire
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.

(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

Upper Guinea: Guinea, Cte dIvoire, 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
Spp
422
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
K.spp.
151
27
64
27
62
11
19
75
All sites
198
26
57
31
58
11
25
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)
11/11/03
4:13 PM
Page 423
Khaya ivorensis
A.Chev.
Meliaceae
Description
Guild: np
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
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, Cte dIvoire, Ghana, Togo
forest, moist semi-deciduous forest. It is common in
Ghana (Hall & Swaine 1981), A Red List species
(Vulnerable).
Habitat
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
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
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
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
>400m
VW
Soil CMK
L
Soil WHC
L
K.spp.
151
27
64
27
62
11
19
75
All sites
198
26
57
31
58
11
25
72
423
11/11/03
4:15 PM
Page 424
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
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
(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
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
Continent: Benin to Uganda (Voorhoeve 1965)

1965)
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

(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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
K.g.
150
21
61
34
57
26
68
All sites
198
26
57
31
58
11
25
72
11/11/03
4:15 PM
Page 425
Lophira alata
Banks ex Gaertn.
Ochnaceae
Description
Regeneration
Guild: pi
8-25 cm), slightly undulate, glossy dark green,
leaves of sapling much larger (up to 10 x 100 cm)
Inflorescence: terminal, unbranched (lax panicle)
Fruit: dry indehiscent (1.2 x 2.8 cm), winged with
unequal sized wings; 1 seed
Other: a tree with a distinctive, modest crown of
tufted leaves. The flush of new leaves is bright red.
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)
evergreen forest (Hall & Swaine 1981). A Red List
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
from October to November
(Voorhoeve 1965)
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
Data sources
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
L.a.
86
12
60
17
15
53
40
44
49
All sites
198
26
57
31
58
11
25
72
425
11/11/03
4:15 PM
Page 426
Lovoa trichilioides
Harms
Meliaceae
Distribution
1965)
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
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
Leaf: alternate, (im)paripinnately compound, 6-10
(sub)opposite leaflets, elliptic to obovate, mesophyll
Inflorescence: (sub)terminal, branched (panicle,
15-40 cm long)
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.
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
>400m
VW
Soil CMK
L
Soil WHC
L
L.t.
115
16
63
12
10
41
53
32
63
All sites
198
26
57
31
58
11
25
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
Phenology
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),
Akachuku (1984), Kyereh et al. (1993), Hawthorne
(1995a), IUCN Red List (2000)
11/11/03
4:15 PM
Page 427
Mammea africana
Sabine
Guttiferae
Description
Guild: sb
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).

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)
1965)
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
M.a.
102
16
59
13
13
31
60
25
66
All sites
198
26
57
31
58
11
25
72
427
11/11/03
4:15 PM
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. 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
Other: the dark foliage and heavy branches make
the crown rather conspicuous. The bark has
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)
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)
Forest type: moist evergreen forest, moist semideciduous forest, dry semi-deciduous forest,
derived savanna-woodland, riverine forest. A Red
List species (lower risk).
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
M.spp.
156
26
59
29
59
12
23
69
All sites
198
26
57
31
58
11
25
72
Data sources
(1966), De Rouw (1991), Kyereh et al. (1993),
Swaine et al. (1997), IUCN Red List (2000)
11/11/03
4:15 PM
Page 429
Nauclea diderrichii
(De Wild. & T.Durand) Merrill
Rubiaceae
Description
Regeneration
Guild: pi
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
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.
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)
1965)
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
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
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
N.d.
145
21
59
10
10
35
59
26
68
All sites
198
26
57
31
58
11
25
72
429
11/11/03
4:15 PM
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
(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
Spp
430
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)

1965)
semi-deciduous forest, secondary forest (Voorhoeve
1965, Hall & Swaine 1981). It is common in Ghana
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
>400m
VW
Soil CMK
L
Soil WHC
L
N.p.
143
31
64
22
66
13
16
76
All sites
198
26
57
31
58
11
25
72
Deciduousness: mostly evergreen, but Taylor

(1960) records that it may be deciduous for a brief
period in the dry season.
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)
11/11/03
4:15 PM
Page 431
Parinari excelsa
Sabine
Chrysobalanaceae
Description
Regeneration
Guild: pi
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
Fruit: fleshy, ovoid (2.8 x 4.3 cm), coriaceous,
rough, covered with numerous small lenticels,
orange-brown or grey when ripe; 1 seed
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.

(cf. Voorhoeve 1965). Regeneration in high forest is
scarce, but common in broken up forest, logging
roads and young secondary forest (Voorhoeve
1965)
Phenology
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)
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)
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
P.spp.
132
18
61
10
11
36
58
29
65
All sites
198
26
57
31
58
11
25
72
431
11/11/03
4:15 PM
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
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
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
Phenology
Deciduousness: deciduous after the rainy season
(Voorhoeve 1965)
Dispersal: probably by mammals (Hawthorne 1995a)
February; fruiting period from February to April
(Voorhoeve 1965)
Distribution
Uses
Continent: West Africa, south of the

Sahara and Central Africa (Voorhoeve 1965)
secondary forest.
A timber species. The fresh seeds are used for

fishing bait (Voorhoeve 1965).
432
Data sources
(1981), Hopkins (1983), Hawthorne (1995a)
11/11/03
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Page 433
Pericopsis elata
(Harms) Van Meeuwen
Leguminosae-Pap.
Description
Guild: np
Leaf: alternate, imparipinnately compound, 9
leaflets, oblong to lanceolate (2.5 x 6.5 cm), entire
Inflorescence: panicle
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
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 (Aubrville 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
Aubrville (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),
433
11/11/03
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Page 434
(Beauv.) Liben
Lecythidaceae
Description
Regeneration
Guild: pi
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.

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
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).
Spp
434
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
P.m.
148
24
62
29
61
23
72
All sites
198
26
57
31
58
11
25
72
Data sources
Taylor (1960), Voorhoeve (1965), Liben (1971), Hall
& Swaine (1981), Hawthorne (1995a)
11/11/03
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Page 435
(Hook.f.) Brenan
Leguminosae-Mim.
Description
Habitat
Guild: np
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
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)
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
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
P.a.
165
25
58
32
58
10
25
68
All sites
198
26
57
31
58
11
25
72
435
11/11/03
4:16 PM
Page 436
(Welw.) Warb.
Myristicaceae
Distribution
Phenology
Continent: Benin to Uganda, Angola (Voorhoeve 1965)

secondary forest (Voorhoeve 1965, Hall & Swaine
1981)
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
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
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).
Spp
436
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
P.a.
170
26
57
31
58
11
24
69
All sites
198
26
57
31
58
11
25
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)
11/11/03
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Page 437
Rhodognaphalon brevicuspe
(Sprague) Roberty
Bombacaceae
Description
Regeneration
Guild: pi
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)
Ghana (Voorhoeve 1965). Common in Ghana (Hall &
Swaine 1981). Red List species (Vulnerable).
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
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
(1967), Hall & Swaine (1981), Hawthorne (1995a),
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).
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
R.b.
119
24
73
27
66
13
82
All sites
198
26
57
31
58
11
25
72
437
11/11/03
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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)
dIvoire, Ghana
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
Leaf: alternate, palmately compound (5 leaflets),
obovate, mesophyll (2.5-12 x 6-20 cm), entire;
stipules large, herbaceous and persistent
Flower: unisexual; corolla white
Spp
438
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
R.h.
149
29
63
25
63
12
15
76
All sites
198
26
57
31
58
11
25
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)
11/11/03
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Page 439
Sacoglottis gabonensis
(Baill.) Urb.
Humiriaceae
Description
Guild: sw
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

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
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
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).
March; fruiting period from September to October
(Voorhoeve 1965)
439
11/11/03
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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
Leaf: alternate in tufts at the end of branchlets,
simple, obovate, notophyll (2.5-4.5 x 5-10 cm),
entire, coriaceous
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
Spp
440

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)

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
>400m
VW
Soil CMK
L
Soil WHC
L
T.i.
127
24
58
11
32
57
11
26
67
All sites
198
26
57
31
58
11
25
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)
11/11/03
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Page 441
Terminalia superba
Engl. & Diels
Combretaceae
Description
Guild: pi
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
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
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 22C is continuous, but periodic
at 27C (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
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
T.s.
160
28
59
28
61
11
21
71
All sites
198
26
57
31
58
11
25
72
441
11/11/03
4:16 PM
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
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
(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,

Bonnhin 2000). It is a good timber species.
Regeneration
Germination is fast, and a large proportion of the
seeds germinate (Bonnhin 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 (Bonnhin
2000). Seedling growth is fairly constant over a
large range of light environments (Swaine et al.
1997).
Distribution
Continent: Benin to Nigeria (Voorhoeve 1965)
semi-deciduous forest. A Red List species
(Endangered).
Spp
442
Uses
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
T.h.
112
13
72
10
35
56
25
70
All sites
198
26
57
31
58
11
25
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), Bonnhin (2000), Hawthorne & Parren
(2000), IUCN Red List (2000)
11/11/03
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Page 443
Triplochiton scleroxylon
K.Schum.
Sterculiaceae
Description
Regeneration
Guild: pi
Max. diameter: > 136 cm (inventory data Ghana)
lobed for 1/3 of the length, lobes broadly ovate to
triangular, entire
(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
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
Dispersal: by wind
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)
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
T.s.
144
31
63
25
63
12
17
76
All sites
198
26
57
31
58
11
25
72
443
11/11/03
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Page 444
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
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
Dispersal: by animals (Alexandre 1977)
fruiting period from August to October (Voorhoeve
1965). In Cte dIvoire, the fruits are produced
rather irregularly, but usually with two peaks in a
year. Only the smaller trees produce fruits.
Description
Guild: sb
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)
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

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)
Spp
444
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
T.a.
99
13
76
28
64
17
78
All sites
198
26
57
31
58
11
25
72
11/11/03
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Page 445
Zanthoxylum gilletii
(De Wild.) P.G.Waterman
Rutaceae
Description
Guild: pi
Leaf: alternate, imparipinnately compound, approx.
13 opposite or sub-opposite leaflets, oblong to
elongate (5 x 14 cm), crenate
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
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
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 Cte dIvoire (Kahn 1982).
Ross (1954), Taylor (1960), Savill & Fox (1967),

Jones (1974), Hall & Swaine (1981), Kahn (1982),
Hawthorne (1995a)
Spp
Altitude
>400m
VW
Soil CMK
L
Soil WHC
L
Z.g.
64
28
66
30
63
84
All sites
198
26
57
31
58
11
25
72
Z
445
446
11/11/03
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Page 446
11
4:18 PM
Page 447
C.C.H. Jongkind
Checklist of Upper Guinea

forest species
1 1
11/11/03
H A P T E R
fc
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
447
fc
11/11/03
4:18 PM
Page 448
Chapter 11. Checklist of Upper Guinea forest species
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 fleurs dAfrique 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 Floras, 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 Musum National
dHistoire 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 Jahrbcher, Stuttgart, Germany
Bot.Helv. Botanica Helvetica, Basel, Switzerland
Bot. Notiser Botanisca Notiser, Stockholm, Sweden
Brittonia, New York Bot. Gardens, USA
Bull.IFAN Bulletin de lInstitut (Franais)
Fondamental dAfrique Noire, Dakar, Senegal
Bull.MNHN Paris Bulletin du Musum National
dHistoire Naturelle, (section B, Adansonia), Paris,
France (sr. 2, 3 & 4)
Bull.NHM Lond. Bulletin of the Natural History
Museum London (Bot. Ser.), UK
Bull.Soc.Bot.Fr. Bulletin Socit Botanique de France,
448
Paris, France
Candollea, Geneve, Switzerland
Fl.Afr.Centr. Flore dAfrique Centrale (Congo K.Rwanda-Burundi), Meise, Belgium
Fl.Cam. Flore du Cameroun, Yaound, Cameroun
Fl.Gabon Flore du Gabon, Paris, France
Fl.Guinea-Bis. Flora da Guin-Bissao, Lisboa, Portugal
Fl. Malesiana Flora Malesiana, Leiden, the Netherlands
Fl.Senegal Flore illustre de Sngal, 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 lInstitut Franais dAfrique
Noire, Paris, France
Mem.ORSTOM Memoires ORSTOM, Paris, France
Mitt.Munch. Mitt. Bot. Staatssammlung Mnchen,
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.
fc
11/11/03
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Page 449
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 Fe (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 (Fe) Copel. (FWTA, Ferns: 62)
- quadrangularis (Fe) 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 Fe (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 sr. 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 sr. 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 sr. 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 sr. 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 & Ak 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 (Linn) Blume (Fl.Zamb. 9, 1: 79)
Sericostachys scandens Gilg & Lopr. (Boissiera 57: 87)
Amaryllidaceae
Crinum jagus (Thomps.) Dandy (Adansonia sr. 2, 20: 184)
- natans Baker (Adansonia sr. 2, 20: 182)
- ornatum (Ait.) Bury (FWTA 3: 134) = C. zeylanicum
- purpurascens Herb. (Adansonia sr. 2, 20: 180)

- zeylanicum (Linn) Linn (Adansonia sr. 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 Linn (FWTA 1: 728)
Trichoscypha albiflora Engl. (FWTA 1: 736) = T. lucens
- arborea (A.Cheval.) A.Cheval. (Adansonia sr. 3, 23: 250)
- atropurpurea Engl. (FWTA 1: 736) = T. mannii
- baldwinii Keay (Adansonia sr. 3, 23: 251)
- barbata Breteler (Adansonia sr. 3, 23: 252)
- beguei Aubrv. & Pellegr. (FWTA 1: 736) = T. bijuga
- bijuga Engl. (Adansonia sr. 3, 23: 252)
- blydeniae Breteler (Adansonia sr. 3, 23: 255)
- cavalliensis Aubrv. & Pellegr. (Adansonia sr. 3, 23: 255)
- chevalieri Aubrv. & Pellegr. (FWTA 1: 736) = T. lucens
- laxissima Breteler (Adansonia sr. 3, 23: 256)
- liberica Engl. (Adansonia sr. 3, 23: 256)
- linderi Breteler (Adansonia sr. 3, 23: 258)
- longifolia (Hook.f.) Engl. (Adansonia sr. 3, 23: 258)
- lucens Oliver (Adansonia sr. 3, 23: 259)
- mannii Hook.f. (Adansonia sr. 3, 23: 261)
- oba Aubrv. & Pellegr. (FWTA 1: 736) = T. lucens
- olodiana Breteler (Adansonia sr. 3, 23: 261)
- smeathmannii Keay (FWTA 1: 736) = T. smythei
- smythei Hutch. & Dalziel (Adansonia sr. 3, 23: 263)
- sp.A. (FWTA 1: 736) = T. barbata
- sp.B. (FWTA 1: 736) = T. barbata
- yapoensis Aubrv. & 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 Aubrv. & 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 sr. 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 & Mendona (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
452
- occidentale Le Thomas (Adansonia sr. 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 Aubrv. (Aubrv. 1959) = Xylopia sp.?
- villosum (Chipp) Aubrv. (Aubrv. 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 Linn (WAUP 2001: 35)
Clitandra cymulosa Benth. (BJBB 58: 159)
Conopharyngia chippii Stapf (Aubrv. 1959) = Tabernaemontana africana
- durissima Stapf (Aubrv. 1959) = Tabernaemontana crassa
- jollyana Stapf (Aubrv. 1959) = Tabernaemontana crassa
- longiflora Stapf (Aubrv. 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. (Aubrv. 1959) = F. africana
Holarrhena africana A.DC. (Aubrv. 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. (Aubrv. 1959) = H. floribunda
- wulfsbergii Stapf (Aubrv. 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 (Aubrv. 1959) =
V. thouarsii
- thouarsii Roem. & Schult. (AUWP 85-3: 39)
Aquifoliaceae
Ilex mitis (Linn) 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 Aubrv. & Pellegr. (FWTA 1: 751)
Polyscias fulva (Hiern) Harms (FWTA 1: 750)
Schefflera barteri (Seem.) Harms (FWTA 1: 751)
Aristolochiaceae
Aristolochia embergeri Nozeran & N.Hall (Adansonia sr. 2, 4: 101)
Pararistolochia flos-avis (A.Cheval.) Hutch. & Dalziel (FWTA 1: 79) = P.
macrocarpa
- goldieana (Hook.f.) Hutch. & Dalziel (Adansonia sr. 2, 17: 484)
- leonensis (Mast.) Hutch. & Dalziel (Adansonia sr. 2, 17: 480)
- macrocarpa (Duch.) Poncy (Adansonia sr. 2, 17: 488)
- mannii (Hook.f.) Keay (Adansonia sr. 2, 17: 478)
- promissa (Mast.) Keay (Adansonia sr. 2, 17: 491)
- zenkeri (Engl.) Hutch. & Dalziel (Adansonia sr. 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 (Linn) 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)
453
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- oculata N.E.Br. (FWTA 2: 96)
Avicenniaceae
Avicennia africana P.Beauv. (FWTA 2: 448) = A. germinans
- germinans (Linn) Linn (Fl.Cam. 19: 60)
- nitida Jacq. (Aubrv. 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-Flix) 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 (Linn) 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) Aubrv. (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) Hall
(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 Hall (Fl.Cam.: 138)
Hippocratea africana Hall (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.Hall) 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 Hall (Fl.Cam.: 212)
- apocynoides (Welw. ex Oliv.) Hall ex J.Ra var. guineensis (Hutch. & Moss)
N.Hall (Fl.Cam.: 204)
- clematoides (Loesener) R.Wilczek (Fl.Cam.: 196)
- ectypetala N.Hall (Mem.IFAN 64: 115)
- iotricha (Loesener) N.Hall (Fl.Cam.: 210)
- rowlandii (Loesener) N.Hall (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.Hall (Fl.Cam.: 190)
Pristimera luteoviridis (Exell) N.Hall var. luteoviridis (Fl.Cam.: 161)
- paniculata (Vahl) N.Hall (Fl.Cam.: 164)
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- plumbea (Blakelock & Wilczek) N.Hall (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.Hall 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.Hall (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.Hall (Mem.IFAN 64: 216)
- lehmbachii Loesener var. leonensis (Hutch. & Moss) N.Hall (Fl.Cam.: 54)
- lehmbachii Loesener var. aurantiaca (N.Hall) N.Hall (Fl.Cam.: 54)
- letestui Pellegr. (Fl.Cam.: 36)
- lomensis Loesener (FWTA 1: 632) = S. stuhlmanniana
- longipes (Oliver) N.Hall var. camerunensis (Loesener) N.Hall (Fl.Cam.: 48)
- miegei N.Hall (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 Hall (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.Hall (Fl.Cam.: 140)
Simirestis atractaspis N.Hall (Bull.MNHN Paris sr. 4, 6: 9)
- dewildemaniana N.Hall (Fl.Cam.: 178)
- staudtii (Loesener) N.Hall (Fl.Cam.: 176)
- tisserantii N.Hall (Fl.Cam.: 180)
Tristemonanthus nigrisilvae (Mem.IFAN 64) N.Hall (N.Hall 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 Linn ssp. atacorensis (A.Chevalier) F.White (BJBB 46: 273)
- icaco Linn 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. (Aubrv. 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 (Linn) 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 Aubrv. ex Keay (FWTA 1: 275)
- hylodendron Mildbr. (FWTA 1: 275)
Quisqualis indica Linn (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 Linn (FWTA 3: 45)
- ascendens Morton (FWTA 3: 47)
- benghalensis Linn 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 (Linn) King & H.Rob. (Candollea 48: 179)
Crassocephalum biafrae (Oliver & Hiern) S.Moore (FWTA 2: 246) = Solanecio b.
Eupatorium odoratum Linn (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 Aubrv. & Pellegr. (Aubrv. 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)
456
Santaloides afzelii (R.Br. ex Planch.) Schellenb. (FWTA 1: 746) = Rourea minor

- gudjuanum (Gilg) Schellenb. (Aubrv. 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 Linn (FWTA 2: 346)
- cairica (Linn) 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 (Linn) 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 Linn (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 (Linn) 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 Aubrv. & 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 Linn (FWTA 3: 152)
- burkilliana J.Mige (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.Mige (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 (Linn) 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. (Aubrv. 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 Grke (BJBB 48: 317)
- heudelotii Hiern (BJBB 48: 322)
- ivorensis Aubrv. & Pellegr. (Aubrv. 1959) = D. mannii
- kamerunensis Grke (BJBB 48: 310)
- kekemi Aubrv. & Pellegr. (Aubrv. 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 Grke (BJBB 48: 300)
457
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- piscatoria Grke (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 Grke (Aubrv. 1959) = D. canaliculata
Maba abyssinica Hiern (Aubrv. 1959) = Diospyros a.
- ferrea (Willd.) Bakhuizen (Aubrv. 1959, 3: 159) = Diospyros f.
- gavi Aubrv. & Pellegr. (Aubrv. 1959) = Diospyros cooperi
- graboensis Aubrv. (Aubrv. 1959) = Diospyros vignei
- soubreana (F.White) Aubrv. (Aubrv. 1959) = Diospyros s.
Erythroxylaceae
Erythroxylum emarginatum Thonn. (FWTA 1: 356)
- mannii Oliver (FWTA 1: 356)
Euphorbiaceae
Acalypha ceraceopunctata Pax (FWTA 1: 409)
- neptunica Mll.Arg. (FWTA 1: 410)
- racemosa Wall. ex Baill. (FWTA 1: 409)
Alchornea cordifolia (Schum. & Thonn.) Muell.Arg. (FWTA 1: 403)
- floribunda Mll.Arg. (FWTA 1: 403)
- hirtella Benth. (FWTA 1: 403)
Amanoa bracteosa Planch. (FWTA 1: 371)
- strobilacea Mll.Arg. (FWTA 1: 371)
Anthostema aubryanum Baill. (FWTA 1: 416)
- senegalense A.Juss. (FWTA 1: 416)
Antidesma laciniatum Mll.Arg. var. laciniatum (FWTA 1: 374)
- laciniatum Mll.Arg. var. membranaceum Mll.Arg. (FWTA 1: 375)
- membranaceum Mll.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 Mll.Arg. (FWTA 1: 370)
- grandis Pierre ex Hutch. (FWTA 1: 370)
- micrantha (Hochst.) Baill. (FWTA 1: 370)
- stenocarpa Mll.Arg. (FWTA 1: 762) = B. micrantha
Cavacoa baldwinii (Keay & Cavaco) J.Lonard (FWTA 1: 762)
Claoxylon hexandrum Mll.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.Lonard (BJBB 30: 438)
Croton aubrevillei J.Lonard (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 Mll.Arg. (FWTA 1: 400)
Crotonogynopsis akeassi J.Lonard (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 Mll.Arg. Pax. & K.Hoffm. (FWTA 1: 382)
- pellegrini Landri (FWTA 1: 381)
- principum (Muell.Arg.) Hutch. (FWTA 1: 381)
458
- singroboensis Ak 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.) Mll.Arg. (BJBB 29: 138)
Gelonium ivorense Aubrv. & Pellegr. (Aubrv. 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 (Aubrv. 1959) = H. heudelotii
- heudelotii Mll.Arg. (FWTA 1: 377)
- lyrata Tulasne (FWTA 1: 377)
Keayodendron bridelioides Landri (BJBB 62: 188)
Macaranga barteri Mll.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 Mll.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.) Mll.Arg. (FWTA 1: 402)
- subulatus Mll.Arg. (FWTA 1: 402)
Manniophyton fulvum Mll.Arg. (FWTA 1: 400)
Mareya micrantha (Benth.) Mll.Arg. (FWTA 1: 404)
Margaritaria discoidea (Baill.) Webster (FTEA: 63)
Martretia quadricornis Beille (BJBB 59: 326)
Mildbraedia paniculata Pax ssp. occidentalis J. Lonard (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.) Mll.Arg. (FWTA 1: 387) = Margaritaria discoidea
- kerstingii Brunel (Willdenowia 15: 251)
- magnificens Brunel & Roux (Willdenowia 11: 82)
- mannianus Mll.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 Mll.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.Lonard (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 (Lonard) 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 (Aubrv. & Pellegr.) J.Lonard (BJBB 28: 449)
- occidentalis (Hoyle) Croizat (BJBB 28: 449)
Tetrorchidium didymostemon (Baill.) Pax & K.Hoffm. (Adansonia sr. 3, 21: 100)
- minus (Prain) Pax & K.Hoffm. (FWTA 1: 414) = T. didymostemon
- oppositifolium (Pax) Pax & K.Hoffm. (Adansonia sr. 3,21: 103)
Thecacoris stenopetala (Muell.Arg.) Muell.Arg. (FWTA 1: 372)
Tragia benthamii Baker (FWTA 1: 412)
- chevalieri Beille (FWTA 1: 412)
- laminularis Mll.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 Linn (FWTA 1: 411)
Uapaca chevalieri Beille (FWTA 1: 390)
- corbisieri De Wild. (Hall & Swaine 1981: 313)
- esculenta A.Cheval. ex Aubrv. & Leandri (FWTA 1: 392) = U. corbisieri
- guineensis Mll.Arg. (FWTA 1: 390)
- heudelotii Baill. (FWTA 1: 390)
- paludosa Aubrv. & 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 sr. 2, 7: 211)
Voyria platypetala Baker (FWTA 2: 302) = V. primuloides
- primuloides Baker (Adansonia sr. 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 Aubrv. & 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 sr. 3, 19: 256)
- brevipes Stapf (Adansonia sr. 3, 19: 256)
- dentata Oliver (Adansonia sr. 3, 19: 257)
- echinata Oliver (Adansonia sr. 3, 19: 257)
- gilgiana Sprague (Adansonia sr. 3, 19: 257)
- glauca (P.Beauv.) Hook.f. (Adansonia sr. 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 (Linn) Desv. (van der Zon 1992: 83)
Coix lacryma-jobi Linn (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 Linn (van der Zon 1992: 38)
Oplismenus burmannii (Retz.) P.Beauv. (van der Zon 1992: 192)
- hirtellus (Linn) P.Beauv. (van der Zon 1992: 193)
Panicum brevifolium Linn (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 Linn (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 Linn f. (FWTA 1: 293)
Vismia guineensis (Linn) 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.Lonard (FWTA 1: 473)
- elongata (Hutch.) J.Lonard (FWTA 1: 473)
- ernae (Dinklage) J. Lonard (FWTA 1: 473)
- explicans (Baill.) J.Lonard (FWTA 1: 473)
- fragrans (Baker f.) Excell & Hillcoat (FWTA 1: 473)
- macrophylla P.Beauv. (FWTA 1: 473)
- sassandraensis Aubrv. & Pellegr. (Boissiera 57: 287)
- vignei (Hoyle) J.Lonard (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 (Linn) Roxb. (FWTA 1: 481)
Cassia alata (Linn) Roxb. (FWTA 1: 452) = Senna a.
- aubrevillei Pellegr. (FWTA 1: 452)
- fikifiki Aubrv. & Pellegr. (Boissiera 57: 289)
- sieberiana DC. (FWTA 1: 452)
- tuhovalyana Ak 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)
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.Lonard (Blumea 12: 220)
- engleri Dinklage & Harms (Blumea 12: 225)
- idae Oldeman, de Wit & J.Lonard (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.Lonard (FWTA 1: 477)
- bilineatum (Hutch. & Dalziel) J.Lonard (FWTA 1: 477)
- ivorense (A.Cheval.) J.Lonard (FWTA 1: 477)
- limba (Scott-Elliot) J.Lonard (FWTA 1: 477)
- obliquum (Stapf) J.Lonard (FWTA 1: 477)
- preussii (Harms) J.Lonard (FWTA 1: 477)
- robynsianum Aubrv. & Pellegr. (Boissiera 57: 294)
- splendidum (Hutch. & Diels) J.Lonard (FWTA 1: 475)
- taiense Aubrv. (Aubrv. 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.Lonard (FWTA 1: 465)
- ehie (A.Cheval.) J.Lonard (FWTA 1: 465)
- leonensis J.Lonard (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 & Mige (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 & Mendona (FWTA 1: 641)
- mangenotiana Mige (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 & Mige) 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 (Grke) J.K.Morton (FWTA 2: 463)
Lauraceae
Beilschmiedia bitehi Aubrv. (Aubrv. 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) =
- 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.Lonard (FWTA 1: 474)
Pellegriniodendron diphyllum (Harms) J.Lonard (FWTA 1: 474)
Plagiosiphon emarginatus (Hutch. & Dalziel) J.Lonard (FWTA 1: 464)
Polystemonanthus dinklagei Harms (FWTA 1: 468)
Senna alata (Linn) Roxb. (Lock 1989: 36)
- tuhovalyana (Ak Assi) Lock (Kew Bull. 43: 341)
Stachyothyrsus stapfiana (A.Cheval.) J.Lonard & 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.Lonard (Voorhoeve 1965: 212)
Toubaouate brevipaniculata (J.Lonard) Aubrv. & Pellegr. (Aubrv. 1959) =
Didelotia b.
Triplisomeris explicans (Baill.) Aubrv. & Pellegr. (Aubrv. 1959) = Anthonotha e.
Leguminosae-Mim.
Acacia ataxacantha DC. (FWTA 1: 499)
- kamerunensis Gand. (Fl.Cam.: 123)
- pennata (Linn) 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 (Linn) 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 Linn (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 sr. 2, 8: 328)
Baphia bancoensis Aubrv. (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 (Linn) 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 (Linn) 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 Linn ssp. occidentalis Verdc. (Kew Bull. 24: 423)
Drepanocarpus lunatus (Linn f.) G.F.W.Mey. (FWTA 1: 519) = Machaerium lunatum
Erythrina addisoniae Hutch. & Dalziel (FWTA 1: 563)
- mildbraedii Harms (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 Ak 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 (Linn) 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 Linn (FWTA 3: 106) = G. superba
- superba Linn (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 sr. 2, 11: 100)
- foliosa Oliver (FWTA 1: 359) = H. afzelii
- planchonii Hook.f. (Adansonia sr. 2, 11: 101)
- platysepala Welw. ex Oliver (Adansonia sr. 2, 11: 99)
- rufipilis A.Cheval. ex Hutch. & Dalziel (Adansonia sr. 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 Linn (FWTA 1: 346)
- tiliaceus Linn (FWTA 1: 345)
- whytei Stapf (FWTA 1: 347)
Thespesia populnea (Linn) 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.) Lonard & 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 Linn (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 sr. 2, 13: 444)
Bourdaria felicis A.Cheval. (FWTA 1: 251) = Cincinnobotrys f.
Calvoa hirsuta Hook.f. (Bull.MNHN Paris sr. 4, 3: 132)
- monticola A.Cheval. ex Hutch. & Dalziel (Bull.MNHN Paris sr. 4, 3: 142)
- trochainii Jacq.-Flix (Bull.MNHN Paris sr. 4, 3: 129)
Cincinnobotrys felicis (A.Cheval.) Jacq.-Flix (Adansonia sr. 2, 16: 377)
Dicellandra barteri Hook.f. var. barteri (Adansonia sr. 2, 14: 86)
Dichaetanthera africana (Hook.f.) Jacq.Flix (Fl.Cam.: 55)
- echinulata (Hook.f.) Jacq.-Flix (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.
- multiflora (Sm.) Triana (Fl.Cam.: 20)
- paucistellata Stapf (FWTA 1: 259) = Melastomastrum afzelii var. p.
- sylvestris Jacq.-Flix (FWTA 1: 256) = Heterotis s.
Guyonia ciliata Hook.f. (Fl.Cam.: 8)
Heterotis antennina (Sm.) Benth. (Adansonia sr. 2, 20: 418)
- entii (J.B.Hall) Jacq.-Flix (Adansonia sr. 2, 20: 419)
- sylvestris (Jacq.-Flix) Jacq.-Flix (Bull.MNHN Paris sr. 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 sr. 3, 17: 61)
- cornifolium (Benth.) Jacq.-Flix (Bull.MNHN Paris sr. 3, 17: 66)
Memecylon afzelii G.Don var. amoenum Jacq.-Flix (Adansonia sr. 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.-Flix (Adansonia sr. 2, 18: 431)
- fasciculare (Planch. ex Benth.) Naud. (FWTA 1: 263) = Warneckea fascicularis
- fleuryi Jacq.-Flix (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 sr. 2, 18: 424)
- membranifolium Hook.f. (FWTA 1: 263) = Warneckea membranifolia
- memecyloides (Benth.) Exell (FWTA 1: 263) = Warneckea m.
- memoratum Jacq.-Flix (Adansonia sr. 2, 18: 416)
- normandii Jacq.-Flix (FWTA 1: 262)
- occultum Jacq.-Flix (Adansonia sr. 2, 18: 412)
- polyanthemos Hook.f. (FWTA 1: 262)
- ramosum Jacq.-Flix (Adansonia sr. 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.-Flix (FWTA 1: 251) = P. kamerunensis
- kamerunensis Gilg (Adansonia sr. 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.-Flix var. fleuryi (Jacq.-Flix) Jacq.-Flix (Adansonia
sr. 2, 18: 226)
- blakeoides (G.Don) Jacq.-Flix var. blakeoides (Adansonia sr. 2, 18: 226)
Tristemma akeassii Jacq.-Flix (Bull.MNHN Paris sr. 2, 28: 168)
- albiflorum (G.Don) Benth. (Bull.MNHN Paris sr. 2, 28: 180)
- coronatum Benth. (Bull.MNHN Paris sr. 2, 28: 195)
- involucratum Benth. (Bull.MNHN Paris sr. 2, 28: 174)
- mauritianum J.-F.Gmelin (Bull.MNHN Paris sr. 2, 28: 152)
Warneckea cinnamomoides (G.Don) Jacq.-Flix (Fl.Cam.: 164)
- fascicularis (Planch ex Benth.) Jacq.-Flix var. fascicularis (Adansonia sr. 2, 19: 266)
- golaensis (Baker f.) Jacq.-Flix (Adansonia sr. 2, 18: 231)

- guineensis (Keay) Jacq.-Flix (Adansonia sr. 2, 18: 235)
- membranifolia (Hook.f.) Jacq.-Flix (Fl.Cam.: 172)
- memecyloides (Benth.) Jacq.-Flix (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)
- thompsonii Sprague & Hutch. (FWTA 1: 707)
Heckeldora latifolia Pierre (see Blumea 22: 491) = Heckeldora staudtii
- mangenotiana Ak 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) Aubrv. (Aubrv. 1959) =
T. ornithothera
- heudelotii Planch. ex Oliver (FWTA 1: 704) = T. monadelpha
- lanata A.Cheval. (FWTA 1: 705) = T. tessmannii
- martineaui Aubrv. & 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 Ak Assi (BJBB 31: 507)
- ghanensis J.B.Hall (Adansonia sr. 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 (Aubrv. 1959) = B. abyssinica
Menispermaceae
Albertisia cordifolia (Mangenot & Mige) 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 & Mige) 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 & Mige (Troupin 1962: 42) = Albertisia
cordifolia
- cuneatum Keay (Troupin 1962: 31) = Albertisia cuneata
- ferrugineum (Diels) Keay (Troupin 1962: 41) = Albertisia ferruginea
- mangenotii Guillaumet & Debray (Adansonia sr. 2, 4: 315) = Albertisia m.
- scandens Mangenot & Mige (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 Mige (Troupin 1962: 171)
- racemifera Miers (Troupin 1962: 168)
Sphenocentrum jollyanum Pierre (BJBB 53: 59)
Stephania dinklagei (Engl.) Diels (Troupin 1962: 258)
463
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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. (Aubrv. 1959) = Trilepisium madagascariense
Cecropia peltata Linn (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.Hall & Ak Assi (Adansonia sr. 2, 7: 390) =
Dorstenia d.
- kameruniana (Engl.) Lanjouw (FWTA 1: 599) = Dorstenia k.
Dorstenia astyanactis Ak Assi (Adansonia sr. 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. (Aubrv. 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 & Lonard (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 Linn (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 sr. 2, 12: 548)
Pycnanthus angolensis (Welw.) Warb. (FWTA 1: 61)
- dinklagei Warb. (FWTA 1: 61)
- kombo Warb. (Aubrv. 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 sr. 2, 10: 377)
- vestita Jacq.-Flix (Adansonia sr. 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 Ak Assi (BJBB 30: 15) = E. gabonensis
- nigerina A.Cheval. ex Hutch. & Dalziel (FWTA 1: 237)
- obanensis Baker (FWTA 1: 238)
- pobeguinii Aubrv. (FWTA 1: 238)
- salacioides Lawson ex Hutch & Dalziel (FWTA 1: 238)
- tabouensis Aubrv. (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 Linn (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.) Ak 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.Lonard var. lucida J.Lonard (FWTA 1: 648) = S. pustulata
- grandifolia Hook.f. ex Benth. (Aubrv. 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. (Aubrv. 1959) = Chionanthus mannii var.
mannii
- mannii Soler. (FWTA 2: 48) = Chionanthus mannii var. mannii
- nilotica Oliver (FWTA 2: 48) = Chionanthus niloticus
Olea capensis Linn ssp. hochstetteri (Baker) Friis & Green (Kew Bull. 41: 36)
- guineensis Hutch. & C.A.Smith (Aubrv. 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 OBrien (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 sr. 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 dAC: 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 sr. 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 sr. 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 sr. 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 sr. 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 sr. 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 (Guine) 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)
466
- 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 (OBrien) 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. Hall (Adansonia sr. 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 sr. 2, 15: 210)
Rangaeris brachyceras (Summerh.) Summerh. (FWTA 3: 266) = Cribbia b.
- longicaudata (Rolfe) Summerh. (Adansonia sr. 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. (Aubrv. 1959) = R. hookeri
Sclerosperma mannii H.Wendl. (FWTA 3: 161)
Pandaceae
Microdesmis keayana J.Lonard (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 (Aubrv. 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 LHerit. (FWTA 1: 143)
Piperaceae
Peperomia fernandopoiana C.DC. (Bot.Jahrb. 93: 113)
- molleri C.DC. (Bot.Jahrb. 9: 106)
- retusa (Linn f.) Dietr. (Bot.Jahrb. 9: 89)
- rotundifolia (Linn) H.B. & K. (Bot.Jahrb. 9: 85)
- tetraphylla (Forster) Hooker & Arnott (Bot.Jahrb. 9: 72)
- vulcanica Baker & Wright (Bot.Jahrb. 9: 102)
Piper capense Linn f. (FWTA 1: 84)
- guineense Schum. & Thonn. (FWTA 1: 84)
- umbellatum Linn (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 sr. 4, 5: 385)
467
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- adamesii (Taylor) Cusset (Bull.MNHN Paris sr. 4, 5: 388)

- bowlingii (J.B.Hall) Cusset (Bull.MNHN Paris sr. 4, 6: 268)
- harrisii Cusset (Bull.MNHN Paris sr. 4, 5: 374)
- ledermannii (Engl.) Cusset (Adansonia sr. 2, 14: 274)
- minutissima Cusset (Bull.MNHN Paris sr. 4, 6: 252)
- mortonii Cusset (Bull.MNHN Paris sr. 4, 5: 368)
- taylorii Cusset (Bull.MNHN Paris sr. 4, 5: 376)
Macropodiella garrettii (Wright) Cusset (Adansonia sr. 2, 17: 302)
- heteromorpha (Baill.) Cusset (Adansonia sr. 2, 17: 298)
- macrothyrsa (Taylor) Cusset (Adansonia sr. 2, 17: 296)
- taylorii Cusset (Adansonia sr. 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 (Aubrv. 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.-Flix (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 Aubrv. & Pellegr. (Bull.Soc.Bot.Fr.105: 34)
- lescotiana J.G.Adam (Bull.IFAN 33: 496)
- nialatou Aubrv. & Pellegr. (FWTA 1: 283)
- paludosa Hutch. & Dalziel ex Jacq.-Flix (FWTA 1: 283)
- sp.B. (FWTA 1: 283) = C. congoensis
Rhizophora harrisonii Leechman (FWTA 1: 285)
- mangle Linn (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
468
(Kew Bull. 55: 77)

- hypoglauca (Welw. ex Hiern) A.Cheval. (FWTA 2: 158) = B. coffeoides ssp.
hypoglauca
- lepidopoda Hutch. & Dalziel (Aubrv. 1959) = B. coffeoides ssp. hypoglauca
Bertiera adamsii (Hepper) N.Hall (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 (Aubrv. 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 Aubrv. & 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 Aubrv. & Pellegr. (Aubrv. 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 Ak 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 Ak 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)
- halli 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. (Aubrv. 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 (Aubrv. 1959) = Agrocoffeopsis afzelii
- maclaudii A.Cheval. (Aubrv. 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. (Aubrv. 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 Aubrv. & Pellegr. (Aubrv. 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 (Linn) I.M.Johnston (FWTA 2: 205)
- rotundifolia A.Cheval. ex Hepper (FWTA 2: 206) = Hymenocoleus rotudifolius
Hallea ciliata (Aubrv. & Pellegr.) Leroy (Adansonia sr. 2, 15: 66) = H. ledermannii
- ledermannii (K.Krause) Verdc. (Kew Bull. 40: 508)
- stipulosa (DC.) Leroy (Adansonia sr. 2, 15: 66)
Heinsia crinita (Afzel.) G.Taylor (FWTA 2: 161)
- pulchella K.Schum. (Aubrv. 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 Aubrv. & Pellegr. (Aubrv. 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 Aubrv. & Pellegr. (FWTA 2: 161) = Hallea ledermannii
- stipulosa (DC.) O.Ktze. (FWTA 2: 161) = Hallea s.
Monosalpinx guillaumetii N.Hall (Adansonia sr. 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 (Pobguin 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.) Aubrv. (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.Hall (Adansonia sr. 2, 5: 97)
- stipulosa (Hutch. & Dalziel) Hepper (FWTA 2: 169) = Poecilocalyx s.
- sylvicola (Hutch. & Dalziel) Bremek. (FWTA 2: 169)
- ziamaeana (Jacq.-Flix) 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 (Aubrv. 1959, 3: 219) =
P. corymbosa corymbosa
469
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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.Hall (Adansonia sr. 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 Ak 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 (Ak 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 (Ak 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 (Aubrv. & 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.) Aubrv. (FWTA 2: 116, 496)
Sericanthe adamii (N.Hall) 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 (Aubrv. & Pellegr.) Robbr. (BJBB 48: 65)
- trilocularis (Scott-Elliot) Robbr. ssp. trilocularis (BJBB 48: 67)
- trilocularis (Scott-Elliot) Robbr. ssp. paroissei (Aubrv. & 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.-Flix) N.Hall (Adansonia sr. 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 sr. 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 (Aubrv. 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 Aubrv. & Pellegr. (BJBB 57: 133)
- macrophylla K.Schum. (BJBB 49: 339)
- okelensis Hiern var. okelensis (BJBB 57: 93)
- pallens Hiern (BJBB 57: 114)
- paroissei Aubrv. & 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 Aubrv. & Pellegr. (FWTA 2: 152) = Sericanthe t.
- trilocularis (Scott-Elliot) Hutch. & Dalziel (FWTA 2: 152) = Sericanthe t.
- vignei Aubrv. & 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 (Aubrv. 1959) = Pauridiantha a.
- hirtellum Benth. (Aubrv. 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 Aubrv. & Pellegr. (FWTA 1: 688) = Vepris t.
Citropsis articulata (W. ex Spr.) Swingle & Kellerman (FWTA 1: 688)
- mirabilis (A.Cheval.) Swingle & Kellerman (Aubrv. 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 (Aubrv. 1959) = Vepris hiernii
- klaineana Pierre (FWTA 1: 689) = Vepris angolensis
Fagara atchoum Ak 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 Ak Assi (BJBB 30: 400)
- parvifoliola A.Cheval. ex Keay (FWTA 1: 685) = Zanthoxylum lemairei
- psammophila Ak 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 & Mendona (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 (Aubrv. & Pellegr.) W.Mziray (Symb.Bot.Ups. 30: 76)
- verdoorniana (Exell & Mendonca) W.Mziray (Symb.Bot.Ups. 30: 76)
Zanthoxylum atchoum (Ak 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 (Ak 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 (Aubrv. 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 (Aubrv. & Pellegr.) Pellegr. (Aubrv. 1959)
- welwitschii (Hiern) Radlk. (FWTA 1: 722)
Cardiospermum grandiflorum Swartz (FWTA 1: 711)
- halicacabum Linn (FWTA 1: 711)
Chytranthus angustifolius Exell (Fl.Cam.: 100)
- atroviolaceus Baker f. ex Hutch. & Dalziel (FWTA 1: 717)
- bracteosus Radlk. (Adansonia sr. 2, 2: 293) = C. angustofolius
- carneus Radlk. (Fl.Cam.: 94)
- cauliflorus (Hutch. & Dalziel) Wickens (Kew Bull. 24: 345)
- longiracemosus Gilg ex Radlk. (Adansonia sr. 2, 2: 299) = C. carneus
- macrobotrys (Gilg) Exell & Mendona (Fl.Cam.: 111)
- mangenotii N.Hall & A.Assi (Adansonia sr. 2, 2: 295) = C. cauliflorus
- setosus Radlk. (Adansonia sr. 2, 2: 297)
- talbotii (Baker) Keay (Fl.Cam.: 108)
- verecundus N.Hall & Ak Assi (Adansonia sr. 2, 2: 297)
- villiger Radlk. (FWTA 1: 717) = C. carneus
Crossonephelis adamii Fouilloy (Adansonia sr. 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 Aubrv. & 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 Linn (FWTA 1: 710)
Placodiscus attenuatus J.B.Hall (Adansonia sr. 2, 20: 290)
- bancoensis Aubrv. & Pellegr. (FWTA 1: 720)
- boya Aubrv. & Pellegr. (FWTA 1: 720)
- bracteosus J.B.Hall (Adansonia sr. 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 sr. 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.) Aubrv. (FWTA 2: 30) = Synsepalum c.
- chevalieri (Engl.) Aubrv. (Aubrv. 1959) = Synsepalum cerasiferum
Aningeria altissima (A.Cheval.) Aubrv. & Pellegr. (FWTA 2: 24) = Pouteria a.
- robusta (A.Cheval.) Aubrv. & Pellegr. (FWTA 2: 24) = Pouteria aningeri
Aubregrinia taiensis (Aubrv. & 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.) Aubrv. & Pellegr. (Aubrville 1959, 3: 108)
Chrysophyllum africanum A.DC. (Bull.MNHN Paris sr. 4, 11: 459.)
- albidum G.Don (FWTA 2: 27)
- azaguieanum Mige (FWTA 2: 27)
- beguei Aubrv. & Pellegr. (FWTA 2: 27)
- delevoyi De Wild. (FWTA 2: 28) = C. africanum
- giganteum A.Cheval. (FWTA 2: 28)
- letestuanum A.Cheval. (Aubrv. 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 Aubrv. & 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. (Aubrv. 1959) = Tieghemella h.
Endotricha taiensis Aubrv. & Pellgr. (Aubrv. 1959) = Aubreginia t.
Englerophytum oblanceolatum (S.Moore) Penn. (Pennington 1991: 252)
- oubanguiense (Aubrv. & Pellegr.) Aubrv. & Pellegr. (BJBB 59: 159)
- aff. oubanguiense (BJBB 59: 160)
Gambeya subnuda (Baker) Pierre (Aubreville) = Chrysophyllum s.
Gluema ivorensis Aubrv. & Pellegr. (FWTA 2: 19)
Inhambanella guereensis (Aubrv. & Pellegr.) Penn. (Pennington 1991: 142)
Ituridendron bequaertii De Wild. (FWTA 2: 18) not part of Omphalocarpus
Kantou guereensis Aubrv. & Pellegr. (FWTA 2: 23) = Inhambanella g.
Lemonniera batesii H.Lecomte (Aubrv. 1959, 3: 116) = Neolemonniera b.
- clitandrifolia H.Lecomte (Aubrv. 1959, 3: 96) = Neolemonniera c.
Malacantha alnifolia (Baker) Pierre (FWTA 2: 24) = Pouteria a.
- heudelotiana Pierre (Aubrv. 1959) = Pouteria alnifolia
Manilkara lacera (Baker) Dubard (Aubrville 1959, 3: 120) = M. obovata
- multinervis (Baker) Dubard (FTEA: 69) = M. obovata
- obovata (Sabine & G.Don) Hemsl. (FWTA 2: 20)
- sylvestris Aubrv. & Pellegr. (Aubrv. 1959, 3: 122) = M. obovata
- welwitschii (Engl.) Dubard (Aubrv. 1959) = M. obovata
Mimusops andongensis Hiern (FWTA 2: 20)
- kummel Bruce ex A. DC. (FWTA 2: 20)
- warneckei Engl. (Aubrv. 1959) = M. andongensis
Neoboivinella glomeruliflora (Hutch. & Dalziel) Aubrv. & Pel (Aubrv. 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 (Aubrv. 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. (Aubrv. 1959) = Synsepalum a.
Synsepalum afzelii (Engl.) Penn. (Pennington 1991: 248)
- aubrevillei (Pellegr.) Aubrv. & 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) Ak Assi & Gautier (Candollea 55: 282)
- revolutum (Baker) Penn. (Pennington 1991: 249)
- tsounkpe Aubrv. & Pellegr. (FWTA 2: 22)
472
Tieghemella heckelii Pierre ex A.Cheval. (FWTA 2: 21)

Vincentella camerounensis Pierre (Aubrv. 1959) = Synsepalum revolutum
- passargei (Engl.) Aubrv. (FWTA 2: 23) = Synsepalum p.
- revoluta (Baker) Pierre (FWTA 2: 23) = Synsepalum revolutum
Scytopetalaceae
Rhaptopetalum beguei Mangenot (361 Adansonia sr. 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 Aubrv. & Pellegr. (FWTA 1: 692)
Hannoa klaineana Pierre & Engl. (FWTA 1: 691)
Harrisonia abyssinica Oliver (FWTA 1: 690)
Mannia simarubopsis Pellegr. (Aubrv. 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 Linn (FWTA 2: 332) = S. erianthum
Sterculiaceae
Byttneria catalpifolia Jacq. ssp. africana (Mast.) Exell & Mendona (FWTA 1: 314)
- guineensis Keay & Milne-Redh. (FWTA 1: 314)
- ivorensis N.Hall (Adansonia sr. 2, 2: 287)
Cola angustifolia K.Schum. (FWTA 1: 329)
- attiensis Aubrv. & 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 Ak 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 (Linn) 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 & Mendona (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 & Mendona) Jongkind to be
published
- malacocarpa Mast. var. malacocarpa (Mast.) Jongkind to be published
- megalocarpa Juss. (FWTA 1: 305)
- mollis Juss. (Adansonia sr. 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 (Aubrv. 1959) = C. aristata
Holoptelea grandis (Hutch.) Mildbr. (FWTA 1: 593)
Trema guineensis (Schum. & Thonn.) Ficalho (FWTA 1: 592) = T. orientalis
- orientalis (Linn) 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 (Linn) 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 (Linn) 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 sr. 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 Grke (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 Grke (FWTA 2: 444)
- polycephalum Baker (FWTA 2: 444)
- sassandrense Jongkind (Syst. Geogr. Pl. 72: 239)
- silvanum Henriq. var. buchholzii (Grke) Verdc. (FTEA: 110)
- sinuatum Hook. (FWTA 2: 444)
- splendens G.Don (FWTA 2: 444)
- thyrsoideum Grke (FWTA 2: 444)
- umbellatum Poir. (FWTA 2: 442)
- violaceum Grke (FWTA 2: 441)
- volubile P.Beauv. (FWTA 2: 444)
Premna angolensis Grke (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 (Aubrv. 1959) = V. ferruginea
- grandifolia Grke (FWTA 2: 446)
- micrantha Grke (FWTA 2: 446)
- oxycuspis Baker (FWTA 2: 446)
- phaeotricha Mildbr. ex Pieper (FWTA 2: 447)
- rivularis Grke (FWTA 2: 446)
- rufa A.Cheval. (Aubrv. 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 (Aubrv. & 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 Tchoum (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 Tchoum (183 Boissiera 58: 183)
- vogelii Hook.f. (FWTA 1: 680) = Cyphostemma v.
Cyphostemma adamii Desc. (Adansonia sr. 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
474
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
476
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
477
478
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C
Appendices
Appendices 1-5
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Page 481
Colour figures Chapter 2: Forest cover changes

in Cte dIvoire and Upper Guinea
4:19 PM
P P E N D I X
Appendix
11/11/03
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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Appendix 1. Colour figures Chapter 2: Forest cover changes in Cte dIvoire and Upper Guinea
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 Cte d'Ivoire,
including Ta 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.
482
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Page 483
483
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Page 484
Figure 2.3 Forest cover in Cte dIvoire 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).
484
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Page 485
Figure 2.4 Forest cover in Cte dIvoire in 1993 (Dao 1999) based on NOAA-AVHRR images. The Ta forest area represents at least 40% of the
total forest area of Cte d'Ivoire. Also the forest area south of Abengourou (the classified forests of Yaya-Bossmati-Mabi) is very important.
485
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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.
486
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(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.
487
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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.
488
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(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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The Hir 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 forts classes 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 Hir (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 fort classe was completely
transformed in oil palm plantations.
The G-Bodinou 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 G-Bodinou fort classe. The landscape
to the north of this forest is made up of
numerous groves. Para-rubber tree and palm
plantations have increased heavily. This fort
classe, along with those of Yapo and Niegr, are
the three last large areas of forest in the South
zone of the country.
The Lovigu 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 forts classes. In 1969,
deforestation already reached the borders of the
forts classes. In 1990, over 40% of the forts
classes of Maf and Lovigu were already
cleared, and in the decade to 2000, most
clearing was done in the Maf fort classe.
This block, along with that of Abi, 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 fort classe 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 Abi block north of the Yapo fort classe.

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 forts classes
(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).
491
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Appendix
Forest sites in Upper Guinea (Chapter 4)
P P E N D I X
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Page 494
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
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
Section 2
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 Nigr transect 1
FC Nigr transect 2
FC Nigr 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
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
9.82
463
2445
54.3
35.5
WE3
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
?
?
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
Cte d'Ivoire
166
158
154
151
150
152
136
137
135
494
Goin-Cav79
HauteDodo
HtSasRec1
HtSasRec2
HtSasRec3
HtSasRec4
NigrRec1
NigrRec2
NigrRec3
11/11/03
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Page 495
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
NigrRec4
NigrRec5
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 Nigr transect 4
FC Nigr 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
Perimtre industriel N
Perimtre industriel SW
Perimtre industriel Centre
Perimtre industriel E
Perimtre 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
495
11/11/03
4:32 PM
Page 496
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
11/11/03
4:32 PM
Page 497
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
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
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
Cte 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
Ta
Ta RvR
213
122
214
220
Tamin
Yapo
Yaya
Zagne
Banco National Park

ICFE Bossmati FC rapp 6
(Lennertz)
IFCE Inv. forets de l'Est
(Laumans)
Bossmati
Dassioko
Djapadji
Haute Dodo
Dogbo
Port Gauthier
Guiroutou
Mabi
Marahue
Monogaga
Near Para village
in Tai National Park
Songan
Perimtre Papetiere sect 1
Monogaga
San Pedro
Nero
Djakoteti
Palabod
Perimtre Papetiere Extension
sect 1 Hana
Perimtre Papetiere Extension
sect 2
Ta National Park
Near Ta village
in Ta National Park
FC Tamin
FC Yapo
FC Yaya
Near Zagne village
in Ta 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
497
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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
Clment & Guinaudeau (1973)
Hawthorne 1995a, 1996, Hawthorne & Abu Juam 1995
De Koning 1983
Kouame et al. chapter 5
Ak Assi 1997
Adou et al. unpubl.data
Kouam 1998b
Jongkind et al. 1999
Van Rompaey 1993
Clment 1973
Ak Assi & Pfeffer 1975
SODEFOR 1986
4:33 PM
Page 499
Appendix
Forest reserves in Upper Guinea
P P E N D I X
11/11/03
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
500
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Page 500
11/11/03
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Page 501
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
Appendix
P P E N D I X
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).
501
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Appendix
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Page 503
Geographical map of West Africa
5
P P E N D I X
503
504
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Page 504
11/21/03
2:22 PM
Page 505
D
Sources &
index
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Authors, photographers and illustrators
Authors
Belign, V. B.P. 708, Yamoussoukro, Cte dIvoire

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 Genve, 1 ch. de
lImpratrice, Case Postale 60, CH-1292 Chambsy/Genve, Switzerland
Dao, H. Universit de Genve, Departement de Geographie, Boulevard du Pont
dArve 40, CH-1211 Genve 4, Switzerland
Gautier, L. Conservatoire et Jardin botaniques de la Ville de Genve, 1 ch. de
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, Universit de Cocody 22, B.P. 582,
Abidjan 22, Cte dIvoire
Kouam, F.N. Laboratoire de Botanique, Universit de Cocody 22, B.P. 582,
Kouassi, K. Laboratoire de Botanique, Universit de Cocody 22, B.P. 582,
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
Siepel, A. Forest Ecology and Forest Management Group, Department of
Spichiger, R. Conservatoire et Jardin botaniques de la Ville de Genve, 1 ch. de
Traor, D. Laboratoire de Botanique, Universit de Cocody 22, B.P. 582,
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
Kouam, 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
518
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Index
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
Bossemati 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
Como 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
Cte 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
Marahou 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
Ta 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
519
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Index
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-Baoul 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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Page 521
Index
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
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
521

L. Poorter, F. Bongers, F.N. Kouamý, W.D. Hawthorne Biodiversity of West African Forests 2004

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L. Poorter, F. Bongers, F.N. Kouamý, W.D. Hawthorne Biodiversity of West African Forests 2004

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22466 layout a-f, 1-2

Biodiversity of West African Forests

22466 layout a-f, 1-2

CABI Publishing is a division of CAB International

Library of Congress Cataloging-in-publication Data

ISBN 0 85199 734 1

22466 layout a-f, 1-2

22466 layout a-f, 1-2

22466 layout a-f, 1-2

West African forests: introduction

Implications for conservation and management

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

Forest cover changes in Cte dIvoire and Upper Guinea

The forest-savanna transition in West Africa

The forests of Upper Guinea: gradients in large species composition

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

Floristic diversity of closed forests in Cte dIvoire

Biodiversity hotspots in West Africa; patterns and causes

M. Holmgren, L. Poorter and A. Siepel

22466 layout a-f, 1-2

Ecological profiles of rare and endemic species

Ecological profiles of large timber species

Checklist of Upper Guinea forest species

Forest sites in Upper Guinea

Forest reserves in Upper Guinea

Hotspots of plant biodiversity in Upper Guinean forests

Geographical map of West Africa

D Sources & index

Authors, photographers and illustrators

22466 layout a-f, 1-2

in West Africa are

The atlas strongly draws on the use of herbarium collections.

22466 layout a-f, 1-2

The following authors contributed to the atlas chapters (in

Horn, Michael Huston, Carel Jongkind, Franois Kouam,

Additional acknowledgements for specific chapters may be

Lourens Poorter, Frans Bongers, Franois Kouam, William Hawthorne

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22466 layout Chapter 1

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

West African forests:

22466 layout Chapter 1

Upper Guinean forests

The rainforests of West Africa have been earmarked as

A distinct biogeographic region

Moroccan Berbers, who referred to the lands south of the

22466 layout Chapter 1

Chapter 1. West African forests: introduction

Gambia river. Another large mangrove area is found south

Figures for animal species are lacking. Myers et al.

Deforestation, threats, and conservation

22466 layout Chapter 1

Chapter 1. West African forests: introduction

Rivers and lakes

Figure 1.2. Map of continental Africa, with potential distribution of

a local uplift of the air, resulting in low pressure near the

22466 layout Chapter 1

Chapter 1. West African forests: introduction

At Cape Palmas, at the Liberian-Ivorian border, the

Figure 1.3 Environmental maps, focusing on the high forest zone of

Granites, gneisses, quartz and schists form the main

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Chapter 1. West African forests: introduction

found, while in valley bottoms most soils are of alluvial