1 2 Natural resources research XX3 Other titles in the series Natural resources research I. A review of the natural reso...
Natural resources research
X X
Other titles in the series Natural resources research I. A review of the natural resources of the African continent II. Bibliography of African hydrology III. Geological m a p of Africa with explanatory note IV. Review of research on latérites V . Functioning of terrestrial ecosystems at the primary production level. Proceedings of the Copenhagen Symposium VI. Aerial surveys and integrated studies. Proceedings of the Toulouse Conference VII. Agroclimatological methods. Proceedings of the Reading Symposium VIII. Proceedings of the Symposium on the Granites of West Africa: Ivory Coast, Nigeria, Cameroon, March 1965 IX. Soil biology. Reviews of research X . Use and conservation of the biosphere X I . Soils and tropical weathering. Proceedings of the Bandung Symposium, 16 to 23 November 1969 XII. Natural resources of humid tropical Asia XIII. C o m p u t e r handling of geographical data X I V . Tropical forest ecosystems. A state-of-knowledge report X V . Review of research on salt-affected soils X V I . Tropical grazing land ecosystems. A state-ofknowledge report XVII. Vegetation m a p of South America: explanatory notes XVIII. Case studies on desertification X I X . Solar electricity
The vegetation of Africa A descriptive memoir to accompany the U n e s c o / A E T F A T / U N S O vegetation map of Africa
by F. White
Unesco
Published in 1983 by the United Nations Educational, Scientific and Cultural Organization, 7 Place de Fontenoy, 75700 Paris Printed by Courvoisier S.A., 2300 L a Chaux-de-Fonds
The designations employed and the presentation of the material in this publication do not imply the expression of any opinion whatsoever on the part of Unesco, A E T F A T or U N S O concerning the legal status of any country or territory, or of its authorities, or concerning the delimitations of the frontiers of any country or territory.
I S B N 92-3-101955-4 © Unesco 1983 Printed in Switzerland
Preface
The n e w vegetation m a p of Africa has been compiled by the Vegetation M a p Committee of the Association pour l'Étude Taxonomique de la Flore de l'Afrique Tropicale ( A E T F A T ) in collaboration with Unesco and the United Nations Sudano-Sahelian Office ( U N S O ) . It comprises three m a p sheets at a scale of 1:5000000, a legend, and the present accompanying memoir. A n earlier A E T F A T Vegetation Map of Africa South of the Sahara was published in 1958 with financial support from Unesco. It soon went out of print, and at the fifth plenary meeting of A E T F A T held in Genoa and Florence in 1963 a small committee was asked to look into the possibility of preparing a new edition. At about this time, Unesco had convened a Standing Committee on Classification and Mapping of Vegetation on a World Basis, and had initiated a programme of mapping the world's vegetation at a scale of 1:5000000. In this connection, A E T F A T was invited to participate in the preparation of a new and more detailed vegetation m a p of the whole of Africa as part of the world series. It was originally intended that all the maps in this series should use a uniform legend and colour scheme, but, because of the complexity of the subject-matter and the diversity of approaches, this objective has not been fully achieved. Hence, the m a p of Africa differs in some important respects from the U n e s c o - F A O Vegetation Map of the Mediterranean Basin and the Unesco Vegetation Map of South America. T h e classification used for the African m a p also diverges in some respects from that recommended in the Unesco publication International Classification and Mapping of Vegetation (Ecology and Conservation Series N o . 6, 1973). It is based almost entirely on physiognomy and floristic composition of the vegetation, and not on climate, although a few comparative climatic terms such as moist and dry are occasionally used in the designation of the mapping units. Otherwise, vegetation and climate are dealt with separately, and separate climatic maps are given in the text for each of the major phytogeographic regions. A feature of the U n e s c o / A E T F A T / U N S O Vegetation Map of Africa is that, in the legend, the mapping units are grouped in the traditional manner according to physiognomy, whereas in the text, here, they are grouped according to the floristic regions in which they occur. There are thus two interconnected classifications, which can be used independently but are fully cross-referenced. The legend permits easy comparison of African
vegetation with that of other continents, whereas in the text it is possible to deal effectively with complicated spatial and dynamic relationships. While the legend of the m a p is composite (English and French), this accompanying memoir, because of its length, has been prepared in separate English and French versions. The memoir aims to provide a succinct though comprehensive account of the vegetation of the African mainland, Madagascar and the other offshore islands. Brief introductory chapters deal with geology, climate, soils, animals, fire, land use and conservation. Their purpose is merely to provide an entrée to the specialist literature and to introduce important topics which recur elsewhere in the main text. The vegetation of the main floristic regions is described individually in twenty-two chapters, which comprise the greater part of the text. For each region the salient features of theflora,geology and climate are also described, and a black-and-white m a p is provided. T h e latter illustrates topographic features mentioned in the text and summarizes the regional climate by means of climatic diagrams. For each of the main vegetation types, references to source materials and other important publications, published photographs and profile diagrams (if available), as well as major synonymy, are given. Publication of the Vegetation Map of Africa forms part of Unesco's long-term programme for the synthesis and diffusion of information on natural resources. The m a p thus complements other maps such as the vegetation maps of the Mediterranean Basin and of South America, the F A O - U n e s c o Soil Map of the World or the small-scale m a p showing the world distribution of arid regions. It is also related to a number of other Unesco initiatives for synthesis of information at regional and international levels in order to promote the integrated management of natural resources. Mention might thus be m a d e of A Review of the Natural Resources of the African Continent (1963), the more recent UnescoU N E P - F A O state-of-knowledge reports on tropical forest ecosystems (1978) and tropical grazing land ecosystems (1979), and a series of national case studies on desertification (1980). Several issues in the M a n and the Biosphere ( M A B ) Technical Notes Series also deal with problems of natural resources and their management in the African region, including a review of ecological approaches to land use in the Sahel ( M A B Technical
Notes 1, 1975), a study on traditional strategies and modern decision-making in the management of natural resources in Africa ( M A B Technical Notes 9, 1978) and an analysis of trends in research and in the application of science and technology for arid-zone development ( M A B Technical Notes 10,1979). The United Nations Sudano-Sahelian Office ( U N S O ) was established by the Secretary-General of the United Nations in 1973, in the aftermath of the severe 1968-73 drought which devastated the economy and social life of the Sahelian region, to initiate and assist in the implementation of the medium- and longterm recovery and rehabilitation programme in the eight countries of the area, namely, Cape Verde, Chad, G a m bia, Mali, Mauritania, Niger, Senegal and Upper Volta. Since that time, U N S O has developed into the principal body and central co-ordinating point of the United Nations system mandated by the General Assembly and other United Nations organs (a) to assist the eight drought-stricken Sahelian countries—members of the Permanent Inter-State Committee for Drought Control in the Sahel (CILSS)—in the implementation of their medium and long-term recovery and rehabilitation programmes; and (b) to act as the arm of the United Nations responsible for assisting, on behalf of the United Nations Environment Programme ( U N E P ) , eighteen countries of the Sudano-Sahelian region in the implementation of the Plan of Action to Combat Desertification, as a joint U N D P / U N E P venture (Djibouti, Ethiopia, Guinea, Guinea-Bissau, Kenya, Nigeria, Somalia, Sudan, Uganda and the United Republic of Cameroon, in addition to the eight countries mentioned above). U N S O ' s work, undertaken in close collaboration with the Sudano-Sahelian countries, C I L S S and the respective United Nations agencies, is focused primarily on: (a) assisting the countries and C I L S S in the planning and programming of priority projects and programmes in the field of drought-related medium- and long-term recovery and rehabilitation activities and desertification control; (b) providing assistance in the mobilisation of the necessary resources for implementing such projects and programmes, either on bilateral or multilateral bases, or by contributions to the United
Nations Trust Fund for Sudano-Sahelian Activities established by the Secretary-General for that purpose; (c) managing the Trust Fund and implementing, from the resources of the Fund, in accordance with the relevant rules and regulations of the United Nations, projects not undertaken bilaterally or multilaterally; and (d) monitoring, reporting and disseminating knowledge on drought-related and desertification-control programmes. A s is shown in Figure 1 (see page 12), the SudanoSahelian region extends over a large portion of Africa. Consequently, it is hoped that the information contained in the Vegetation Map of Africa and in the accompanying memoir will provide a synthesis of knowledge on African vegetation that will be useful as a source of reference in land-use planning as well as in training for the purposes of drought recovery and rehabilitation and of desertification control. The task of preparing the m a p and memoir has been a very long and complex one, and Unesco expresses its sincere thanks to the A E T F A T Vegetation M a p Committee in seeing the work through to fruition. The committee was made up of the following specialists: A . Aubreville, L . A . G . Barbosa, L . E . Codd, P. Duvigneaud, H . Gaussen, R . E . G . Pichi-Sermolli, H . Wild and F. White (Secretary). In publishing the new Vegetation Map of Africa, Unesco is especially grateful to M r Frank White of Oxford University (United Kingdom), w h o compiled the vegetation m a p on behalf of the A E T F A T Vegetation M a p Committee and w h o is the author of the present memoir. The views expressed herein are those of the author and are not necessarily shared by Unesco and
by UNSO. Unesco also thanks the Oxford University Press for the preparation of successive proofs of the m a p . In finalizing the m a p and memoir, an attempt has been m a d e to use up-to-date forms of geographic names. However, the designations employed and the delimitations of frontiers on the m a p and in the accompanying text do not imply the expression of any opinion whatsoever on the part of Unesco or U N S O concerning the legal or constitutional status of any country, territory, city or area or of its authorities, or concerning the delimitation of its frontiers or boundaries.
Contents
Introduction
9
Acknowledgements
13
List of former n a m e s of countries 1S Part O n e
Environment, land use and conservation 1. Geology and physiography 19 2. Climate and plant growth 2 3 3. SoUs 2 6 4. Animals 2 9 5. Fire, land use and conservation 3 2 Part T w o Regional framework, classification and mapping units Introduction 3 7 6. Regional framework 39 7. Classification 4 4 8. M a p p i n g units 56 Part Three Vegetation of the floristic regions Introduction 6 9 THE AFRICAN MAINLAND
I. II. III.
T h e Guineo-Congolian regional centre of e n d e m i s m T h e Zambezian regional centre of e n d e m i s m 86 T h e Sudanian regional centre of e n d e m i s m 102
IV.
The Somalia-Masai regional centre of endemism
V.
T h e C a p e regional centre of e n d e m i s m 131
VI.
The Karoo-Namib regional centre of endemism
71
110 136
VII. T h e Mediterranean regional centre of e n d e m i s m 146 VIII/IX. T h e Afromontane archipelago-like regional centre of e n d e m i s m and the Afroalpine archipelago-like region of extreme floristic impoverishment 161 X. T h e Guinea-Congolia/Zambezia regional transition zone 170 XI.
The Guinea-Congolia/Sudania regional transition zone
XII. XIII.
T h e Lake Victoria regional mosaic 179 T h e Zanzibar-Inhambane regional mosaic 1 8 4
XIV. The Kalahari-Highveld regional transition zone 190 XV. The Tongaland-Pondoland regional mosaic 197 XVI. The Sahel regional transition zone 203 XVII. The Sahara regional transition zone 216 XVIII. The Mediterranean/Sahara regional transition zone 225 MADAGASCAR AND OTHER OFFSHORE ISLANDS XIX. The East Malagasy regional centre of endemism 234
XX.
T h e W e s t Malagasy regional centre of e n d e m i s m 2 4 0
XXI.
Other offshore islands
244
175
AZONAL VEGETATION
XXII.
Mangrove, halophytic and fresh-water s w a m p vegetation
260
Glossary and index of vernacular names of vegetation types and habitat Geographical bibliography
271
Alphabetical bibliography
275
Index of plant names
325
269
Introduction
T h e n e w Vegetation Map of Africa and its accompanying text is the fruit of s o m e fifteen years of co-operation between Unesco a n d A E T F A T (Association pour l'Étude Taxonomique de la Flore de l'Afrique Tropicale). In 1965 the A E T F A T Vegetation M a p Committee, consisting of A . Aubrévüle, L . A . G . Barbosa, L . E . C o d d , P. Duvigneaud, R . E . G . Pichi-Sermolli, H . Wild and F . White (Secretary), to which the late H . Gaussen w a s subsequently co-opted, w a s asked to collaborate with U n e s c o in the preparation of a n e w vegetation m a p of Africa as part of the latter's p r o g r a m m e of mapping the world's vegetation at a scale of 1:5000000. T h e materials used in compiling this m a p turned out to b e exceedingly diverse. Those used for the first draft included: 1. 2.
3.
4. 5. 6.
Original contributions by Duvigneaud, Pichi-Sermolli and Gaussen for Zaire, the Ethiopian region, and the Maghreb and Madagascar respectively. Large-scale maps which were being prepared for independent publication elsewhere by Wild & Barbosa for the Flora Zambesiaca region (1:2500000) and by Barbosa for Angola (1:2500000). The remarkably detailed and accurate map of the Veld types of South Africa (1:1500000) by Aco*cks. Codd advised on the adaptation and simplification of this work for the present purpose. Several published and unpublished maps of parts of francophone Africa communicated by Aubrévüle. For much of the remainder of Africa, a large number of vegetation maps at various scales, which had been prepared for a wide variety of purposes. For the few parts of Africa without vegetation maps of any description, correspondence with a host of local specialists, many of w h o m are members of A E T F A T , has supplied the missing information.
I have been responsible for attempting to standardize the source materials and weave them into a coherent whole. In doing this I have continued to receive the unstinted help of the committee m e m b e r s and m a n y others, but I must bear full responsibility for the final presentation and the imperfections which remain. T h e m a p and the descriptive m e m o i r cover not only the whole of Africa and the large island of Madagascar, but also all ecologically important islands in the eastern South Atlantic and western Indian Oceans, though space could only be found for a very brief treatment.
The purpose of the m a p is not to provide detailed information o n any particular area for the benefit of residents in that area, since that information is usually available locally in published or unpublished form and is also inappropriate o n m a p s of this scale. Rather, the purpose is to indicate to local residents in broad terms the manner in which the main features of their local vegetation can be related to the main features of African vegetation as a whole. Another major objective is to provide a framework o n a continental scale within which m o r e detailed local studies can b e conducted a n d compared. Inevitably, in simplifying larger-scale m a p s arbitrary decisions cannot be avoided and deliberate omissions must be m a d e . This should be borne in m i n d by users of the m a p , especially those seeking local detail. It is for considerations such as these that s o m e features of altitudinal zonation have been deliberately suppressed. It would have been cartographically possible to have s h o w n a m o r e complete and accurate zonation but the small gain in factual content would not have been commensurate with the effort and costs involved. A few years elapsed between the completion of the first draft of the m a p and its publication. This time w a s put to good use since it provided m a n y opportunities for testing the accuracy of the m a p . This w a s done in different ways, as follows: 1.
2.
3.
Several ecologists, travelling extensively in Africa, have been able to check the map against vegetation on the ground. I, myself, in the course of three journeys in East, Central and Southern Africa, undertaken for other purposes, have been able to check the accuracy of several pattern lines. The Maghreb was also specifically visited to check the m a p and collect information for the accompanying text. Earlier drafts of the m a p were exhibited at the plenary congresses of A E T F A T held in Munich in 1970 and Geneva in 1974, when members of A E T F A T , whose collective experience, which embraces the whole of Africa, were invited to comment on the map. By this means several inaccuracies were avoided. For those parts of Africa I have not visited a collection of vegetation photographs, published and unpublished, was assembled in order to check whether the physiognomy they portray agrees with that one would expect from the map.
All this checking has led to s o m e significant corrections, but for m u c h of Africa the accuracy of the m a p , within the objectives set out above, w a s confirmed. For a few
10
The vegetation of Africa
(relatively) parts of Africa, there is a total dearth of information. For other parts, even where the original documentation was sparse, subsequent work has usually confirmed the validity of the general features shown on the m a p , though inevitably some local detail was shown to be inaccurate. This reflects, at least in part, inaccuracies in the base m a p s used at different stages in the compilation. The relationship between the broad framework attempted in the U n e s c o / A E T F A T / U N S O Vegetation Map and more detailed studies can be illustrated by reference to the vegetation of Marsabit District in Kenya, which I visited in 1980, long after the m a p was completed. T h e area under consideration occupies approximately 3 x 3 . 5 c m of the Unesco m a p , which shows a pattern of islands of undifferentiated Afromontane communities and degraded evergreen bushland arising from a matrix of lowland deciduous bushland and semi-desert shrubland and grassland. T h e m u c h more detailed m a p of Herlocker (1979a) at a scale ten times greater (area of m a p 100 times larger) shows nine primary vegetation types which can readily be interpreted in terms of the Unesco framework, even though the criteria for distinguishing the major types do not always coincide. The classification of vegetation used in the present work differs in some respects from those in general use. This, in part, is due to the fact that the flora and vegetation of Africa are now better k n o w n than those of most other parts of the tropics. For Africa a new synthesis n o w seems appropriate. T h e principles underlying the present approach are discussed in Chapters 6 and 7. They can be briefly summarized as follows: 1. Vegetation, in the first instance, should be classified without reference to the physical environment, including climate, or to animals. The extent to which environmental factors and the associated fauna can be used to diagnose vegetation types should be evaluated independently. In the classification adopted here climatic terms are occasionally used, but only as a convenient nomenclatural shorthand for important physiognomic and floristic differences which it would be impossible to designate concisely in purely physiognomic or floristic terms. 2. The physiognomic features which figure prominently in conventional classifications were found to be inadequate. 3. A chorological system based o n the patterns of geographical distribution shown by entire floras, as revealed by samples, was found to provide both the basis of an objective framework within which the vegetation of Africa can be described and compared, and an indirect shorthand method of expressing the entire physiognomy of regional vegetation types, not merely a few selected features. 4. A s to nomenclature, it was found that sufficient English terms were available to cover the main physiognomic types, and no difficulty was experienced in finding French equivalents. It was thus possible to avoid the use of imported vernacular names of equivocal application such as savanna and steppe for classificatory units of high rank. By contrast, indigenous, African vernacular names for local variants of the major physiognomic types have proved most valuable.
For some parts of Africa arichvocabulary is available and several terms such as 'muhulu', 'mopane', 'miombo' have been taken up and precisely applied by local botanists. Relatively few such terms have been used in a formal sense in the present work, although the potential application of others is indicated in the text. The fifty or so terms m e n tioned are included in a combined glossary/index at the end of the work. Because the classification is based solely on the plants themselves, sometimes supplemented by features of the environment which can be seen, such as standing water or outcrops of rock, the colour scheme adopted for the m a p does not consciously embody information on climate. The colours chosen were determined primarily by economy and clarity of presentation of information on the vegetation. In order to maintain continuity with the first A E T F A T m a p , the Vegetation Map of Africa South of the Tropic of Cancer (Keay, 1959ft,), which is well known, similar colours have been used for the main regional vegetation types. Since the colours of the earlier m a p were chosen to reflect the principles of Gaussen's (1955) system of climato-ecological cartography, this new m a p inevitably embodies some climatic information. But this is accidental. In m y opinion, in our present state of knowledge, it is preferable to m a p climate and vegetation separately. With this in mind, in the present work an attempt has been m a d e to characterize the twenty regional phytochoria by means of selected climatic diagrams. T h e results are displayed on seventeen pictorialized climatic m a p s (Figs. 5-8 and 11-23) (not twenty, since some figures include more than one phytochorion). These m a p s not only summarize a great deal of climatic information, but also allow visual comparison of the main climatic features of the different phytochoria. They also show all the main topographical features mentioned in the text. Within the constraints imposed by economy and continuity with the past, the colours of the mapping units have been chosen to bring out the relationships of the latter. M u c h of the vegetation of Africa is transitional and this is shown by using 'pyjama' stripes. A white background to stripes of colour indicates landscapes which are largely anthropic. T h e m a p was designed so that, when viewed from a distance, it shows features of regional extent. Increasingly closer inspection should reveal detail of increasingly local significance, but for ultimate detail the user should refer to the text. For reasons given in Part T w o , the one hundred cartographic units shown on the m a p are described in the text under the twenty regional phytochoria (eighteen on the African mainland, two on Madagascar) in which they occur. T h e outlines of these phytochoria are also boldly indicated on the m a p . T o a considerable extent they coincide with the regional vegetation types, although they were discovered independently.
Introduction
The main aim of the text is to describe salient features of the vegetation. The original draft was twice as long as this published version, so, for reasons of econo m y , m u c h detail has been omitted. Similarly little room could be found for detailed discussion of the influence of climate, geology, soils, fire, animals and m a n on vegetation. W h e r e such influences are particularly striking they are mentioned in the text. Otherwise, detail must be sought in the works referred to in the five introductory chapters dealing with these topics. The vegetation types are described as concisely as is compatible with their complexity. For each type the principal sources of information, both published and unpublished, are cited, as are references to characteristic photographs and profile diagrams. It is hoped that this information will, at least in part, compensate for the fact that, diagrams and maps excepted, this work lacks illustrations. In order to avoid imprecise generalizations, I have tried to characterize the individual vegetation types by describing one or more concrete examples. Wherever possible, I have chosen examples I a m familiar with in the field, or those described by authors w h o were available for discussion of their work, either orally or by correspondence. Inevitably, because of the great area covered and the diversity of vegetation types, it has not normally been possible to describe detailed patterns of vegetation in relation to environmental factors and h u m a n interference. There are, however, a few important exceptions. This is chiefly because U N S O expressed a wish that such information should be given for the drier parts of Africa, which suffered from the recent Great Sahelian Drought. This provided an opportunity to include detailed accounts of selected areas of outstanding ecological importance in relation to h u m a n needs. T h e examples chosen are the Jebel Marra area and part of Kordofan Province, Sudan Republic in the Sahel Region, and the Serengeti ecosystem, and part of Marsabit District, Kenya, in the Somalia-Masai Region. They clearly illustrate the ways in which the results of such local studies can be accommodated in a more general framework, such as that which is the subject of the present work, and also demonstrate the fundamental importance of the detailed study and mapping of vegetation in land-use planning. In addition to the detailed studies referred to above, other investigations inspired b y the Great Sahelian Drought have resulted in an extensive, more general, literature on the ecology of the Arid Zone (Anon, 1977; Bartha, 1970; Breman & Cissé, 1977; Brown, 1971; Cloudsley-Thompson, 1974; Curry-Lindahl, 1974; Dalby & Harrison-Church, 1973; Dalby et al, 1977; D e Leeuw, 1965; F A O , 1977; Galláis, 1975; Konczacki, 1978; Lamprey, 1975, 1978; Lewis, 1975; M o n o d , 1975; Petrides, 1974; Swift, 1973; and Unesco, 1975). It has not been possible to summarize this in detail, though it has been taken into account in expanding the chapters on the Somalia-Masai and Sahelian regions.
11
The position of the Sahel and the four study areas mentioned above are shown on Figure 1. In attempting to characterize the different vegetation types floristically, economy has been exercised in the selection of representative species. In other words equivalents of the 'zone fossils' of geologists have been sought. Even so, it has been necessary to mention some 3000 species in the text. They are included in an index to botanical names together with those major synonyms that feature prominently in the ecological literature. Hence the index provides a means of obtaining both new and previously published information on the autecology of the majority of important plant species in Africa. A serious attempt has been m a d e to ensure that the names used are those which satisfy the requirements of the International Code of Botanical Nomenclature and that the citation of authors is correct. In this connection I have received m u c h help from the Director of the Royal Botanic Gardens, K e w , and his staff, but for some parts of Africa, particularly North Africa, the relevant information is not readily available and some further checking is required. T h e names of countries are based upon United Nations official practice. There are a few exceptions. Thus, for the United Republic of Cameroon, the simple French title of 'Cameroun' is used throughout the publication, in order to retain consistency with the names of geographic locations such as M o u n t Cameroun and the Cameroun highlands. For reasons of brevity, the D e m o cratic Republic of the Sudan is referred to either as Sudan or as Sudan Republic when the use of term 'Sudan' could be taken to m e a n either the country or the centre of endemism of the same name. Similarly, 'Guinea' and 'Guinea Republic' are used as synonyms for the Revolutionary People's Republic of Guinea. The Socialist People's Libyan Arab Jamahiriya is referred to as Libya or the Libyan Arab Jamahiriya. Generally, the English-language version of a country's title is used in the text of the memoir (e.g. Ivory Coast), though the French title (Côte d'Ivoire) m a y be used on the m a p and climate diagrams, given that a single version of the m a p and figures has been prepared to accompany the two language versions of this descriptive memoir. Finally, some older administrative names have been maintained because they feature prominently in the botanical literature. The bibliography of some 2400 items aims to be comprehensive rather than complete, though it is unlikely that m a n y important works have been left out. Apart from a few recent publications, most of the works referred to were used in writing this book, and all but a small minority of the cited references were personally checked by the author in the course of completing the manuscript. For ease of reference a second bibliography is provided in which the publications are arranged geographically. F. White
The vegetation of Africa
12
20* -7
10 e 1
0° I
10* 1
20* 1
30* 1
40* <
50° r-
FIG. 1. M a p of Africa showing names of countries and position of the Sahel regional transition zone, the Somalia-Masai regional centre of endemism and four Sahel-type study areas described in the text: (A) Jebel Marra (Chapter X V I ) ; (B) Kordofan (Chapter XVI); (C) South-Western Marsabit District (Chapter IV); (D) The greater Serengeti region (Chapter IV). The following countries are members of the Plan of Action to Combat Desertification: Cape Verde, Chad, Djibouti, Ethiopia, Gambia, Guinea, Guinea-Bissau, Kenya, Mali, Mauritania, Niger, Nigeria, Senegal, Somalia, Sudan, Uganda, United Republic of Cameroon and Upper Volta
Acknowledgements
M a n y people have contributed to the successful completion of this work. It is a pleasure to record their kind help. In addition to the members of the A E T F A T Vegetation M a p Committee and others mentioned in the Introduction, the following either read early drafts or provided information: J. P. H . Aco*cks (South Africa), E . J. Adjanohoun (Benin), L . Aké Assi (Ivory Coast), G . Aymonin (Madagascar), P. Bamps (Zaire and general), J. P. M . Brenan (general), J. F. M . Cannon (general), J. D . Chapman (Malawi, Nigeria), W . D . Clayton (grasslands), M . J. Coe (animals), K . G . Cox (geology), D . Edwards (South Africa), C . Evrard (Zaire), D . B . Fanshawe (Zambia), I. Friis (Ethiopia), M . G . Gilbert (Ethiopia, Kenya), J. B . Gillett (Ethiopia, Kenya), the late P. E . Glover (Kenya), the late P. J. Greenway (Kenya), J. B . Hall (Ghana), A . J. HallMartin (Malawi), O . Hedberg (Afroalpine vegetation), C . F . Hemming (arid zones), C . J. Humphries (Canaries), P. James (Ascension), C . Jeffrey (general), E . W . Jones (Nigeria), D . J. B . Killick (South Africa), F. J. Kruger (Cape), H . F. Lamprey (Serengeti), R . M . Lawton (Nigeria, Zambia), J. P. Lebrun (Sahel), O . Leistner (Kalahari), J. Léonard (Sahara), R . Letouzey (Cameroun, general), A . Le Thomas (Gabon), J. Lewalle (Burundi, Maghreb), L . Leyton (Welwitschia), G . L . Lucas (general), D . J. Mabberley (general), W . S. McKerrow (geology), F . Malaisse (Zaire), W . Marais (Mascarenes), E . J. Mendes (general), H . Merxmüller (Namibia), E . J. Moll (South Africa), T . M o n o d (Sahel), J. K . Morton (Ghana, Sierra Leone), R . M . Polhill (Kenya), D . J. Pratt (general), the late J. Procter (Tanzania), P. Quézel (Sahara), A . Radcliffe-Smith (Socotra), the late J. Raynal (general), A . Raynal-Roques (general), S. A . Renvoize (Aldabra), E . R . C . Reynolds (climate and plant growth), W . A . Rodgers (Tanzania), R . Rose Innes (Ghana), J. H . Ross (South Africa), R . Schnell (West Africa), P. J. Stewart (Maghreb), P. Sunding
(Mascarenes), M . D . Swaine (Ghana), J. J. Symoens (Zaire), T . J. Synnott (general), H . C . Taylor (South Africa), B. Verdcourt (East Africa), the late D . F. VeseyFitzGerald (Tanzania), R . Webster (soils), G . E . Wickens (Sudan, general), M . J. A . Werger (South Africa). In the early stages of this work, R . W . J. Keay kindly made his considerable experience of African vegetation available and gave generously of his time. Assistance in the field was provided by: W . R . Bainbridge (Natal), the Chief Conservator of Forests, Kenya, L. E . Codd and B . de Winter (South Africa), R . B . D r u m m o n d (Zimbabwe), J. B . Gillett (Kenya), D . Herlocker (Kenya), Christine Kabuye (East Africa), J.O. Kokwaro (Kenya), J. Kornaá (Zambia), F.J. Kruger (Cape), H . F. Lamprey (Kenya), J. Lewalle (Morocco), E . J. Moll (Natal), T . Müller (Zimbabwe), J. C . Scheepers (Transvaal), John and Lucie Tanner (Tanzania) and H . C . Taylor (Cape). Three successive Professors of Forest Science in the University of Oxford, M . V . Laurie, J. L . Harley and M . E . D . Poore, have freely made the facilities of their Department available for the execution of this work. A special debt of gratitude is owed to Ernie Hemmings, Librarian in the Department of Forestry, and his staff for the trouble they have taken in obtaining photocopies of rare literature and for other bibliographic assistance. M u c h help was provided by the Assistants in the Forest Herbarium, notably Rosemary Wise, who prepared the text figures, Frances Bennett and Helen Hopkins, who assisted in the compilation of the m a p , Serena Marner and Michael Wilkinson, w h o have helped with the bibliography and index respectively, and Cynthia Styles, who typed the text. I a m also deeply indebted to John Callow and his colleagues at the Oxford University Press for their advice on cartographic matters and their patience in handling a project which extended over several years.
F.W.
List of former names of countries
List of former names of countries which appear in the cited botanical literature. Old name Abyssinia Anglo-Egyptian Sudan Basutoland Bechuanaland Protectorate Belgian Congo British Somaliland Cameroons Dahomey Eritrea Fernando Po French Guinea French Somaliland French Sudan Gold Coast
New name Ethiopia Sudan Lesotho Botswana Zaire Somalia (part) United Republic of Cameroon or Cameroun Benin Ethiopia (part) Bioko Guinea Djibouti Mali Ghana
Old name Italian Somaliland Madagascar Middle Congo Northern Rhodesia Nyasaland Rhodesia Rio Muni Ruanda Southern Rhodesia South-West Africa Tanganyika Territory Ubangi-Shari Union of South Africa Urundi Zululand
New name Somalia (part) Malagasy Republic Congo Zambia Malawi Zimbabwe Equatorial Guinea Rwanda Zimbabwe Namibia Tanzania
Central African Republic Republic of South Africa Burundi KwaZulu (part)
Part O n e
Environment, land use and conservation
20'
10°
0'
10°
FIG. 2. Physical features of Africa (after Grove, 1978)
20°
30°
40°
SO
i
Geology and physiography
Africa is the second largest continent. It is unique in that, with the exception of the Atlas mountain system in the north-west and the Cape ranges in the south, it consists of a continuous crystalline shield which is exposed over extensive areas. In places, unaltered sedimentary rocks deposited on a metamorphic basem*nt complex are largely of Precambrian age. In the Sahara and the Kalahari region aeolian sands conceal the older rocks over large surfaces. A general account of the geology of Africa has been published by Furon (1963, 1968), w h o also wrote the explanatory note to the Unesco/ Asga geological m a p of Africa (Furon & Lombard, 1964). The latter is extremely brief. A somewhat longer treatment accompanies the F A O - U n e s c o soil m a p of the world ( F A O - U n e s c o , 1977). Clear summaries of the main features of African geology are few. The following pages rely heavily on that of Grove (1978). Important regional works include those by Gray (éd., 1971) on the Libyan Arab Jamahiriya, by Whiteman (1971) on the Sudan Republic, by Cahen (1954) on Zaire, by Haughton (1963, 1969) on the stratigraphy of Africa south of the Sahara and on the geological history of southern Africa, by D u Toit (1954) on South African geology, and by King (1967a, 1967b, 1978) on South African scenery and geomorphology. The physiography and geology of East Africa are briefly summarized by Saggerson (1962a, 1962ft, 1972). Màckel (1974) has described the geomorphology of the shallow streamless depressions (dambos) at the heads of drainage systems on the Great African Plateau in south-central Africa. A line drawn across the m a p of Africa from Angola to western Ethiopia divides the continent into high and low parts (Fig. 2). L o w Africa, in the north-west, is m a d e up of sedimentary basins and upland plains mostly between 150 and 600 m above sea level, comprising the Sahara and the catchments of the lower Nile, Senegal, Niger, Chad and Zairerivers.Landrisingabove 1000 m is confined mainly to: the Atlas mountains in the Maghreb; the Sanaran massifs Ahaggar and Tibesti; Jebel Marra in the Sudan Republic; the headwaters of the Niger; the Jos Plateau in Nigeria; and the Cameroun highlands. Nearly all of High Africa, to the south and east,risesabove 1000 m , with the exceptions of Somalia, the broad lowlands of Mozambique and relatively narrow coastal plains and valley strips elsewhere. Even the Kalahari basin is about 1000 m above the sea, and in
east Africa the surface of Lake Victoria stands 1130 m above sea-level. The rocks of Africa span a time period of 3 500 million years and the geological structure of the continent is complicated. There are three great areas where the rocks were affected by mountain building more than 1500 million years ago that have not been disturbed by folding since. They occupy the western lobe of Africa, the ZaireAngola region and the Zimbabwe-Transvaal-Orange Free State area, and are called older cratons. The areas between the older cratons have been affected by m o u n tain building within the last 1200 million years and are k n o w n as the younger orogens. A m o n g those minerals which are associated with vegetation anomalies, the principal deposits of chromium and asbestos are found in the older cratons, whereas most of Africa's copper, lead, zinc, and cobalt are contained in the younger orogens. The ancient rocks of the older cratons and younger orogens, which are mostly of Precambrian age, underlie the whole area, but they are widely buried by younger sedimentary rocks and wind-blown sand. They outcrop in watershed areas and escarpmentsflankingthe Niger basin, and form the rugged massifs of Air and Ahaggar as well as the dissected plateaux of the Guinea highlands, and Jos in Nigeria. They build the Cameroun highlands and all the western rim of the continent from the Crystal Mountains to the Orange River in the south. Granites, gneisses and schists, exposed in hill ranges or masked by the products of long-continued weathering, form long watersheds between the Nile, Zaire and Chad basins. In East Africa they appear more extensively than elsewhere, building m u c h of the high country between the Transvaal in the south and the Red Sea Hills of Egypt in the north. The oldest of the crystalline rocks are intensely folded schists and banded gneisses which are resistant to erosion but generally less so than the granites intruded into them. Over wide areas these old rocks are characterized by extensive, gently sloping surfaces, covered with a deeply weathered layer or comparatively recently transported material which conceals the solid rocks. The granites and some of the gneisses form rugged hills or dome-shaped inselbergs whichrisesharply from the surrounding plains, whereas quartz dykes stand up as long narrow ridges. C o m m o n l y it is the rocks in the vicinity of igneous intrusions that have been mineralized.
20
Environment, land use and conservation
The youngest of the Precambrian rocks have not been strongly folded in most areas. They include the gold-bearing quartzites of the Rand, much-hardened sandstones, of which the older cratonic rocks were the source. F r o m the Lower Palaeozoic period until the Jurassic, Africa, together with South America, Antarctica, M a d a gascar, India and Australia, was part of a southern land mass, Gondwanaland. Before its break up it was surrounded by an ocean in which great layers of sediments accumulated. They n o w form sandstones, shales, limestone and dolomites in the Maghreb, the western Sahara and the Cape. S o m e of these rocks are very resistant to erosion, notably the tough sandstones of the Cape series which form Table Mountain behind Cape T o w n . The sands and clays accumulating on the continental margins and in the oceans around Gondwanaland were derived from the erosion of the surface of the supercontinent. In Ordovician times the seas locally invaded the interior and laid d o w n shales and sandstones which n o w form escarpments rising from the desert plains at the margins of Tibesti and Tassili des Ajjers. Resting on these marine rocks are tillites deposited by the continental glaciers that, 450 million years ago, occupied m u c h of what is n o w the north-western Sahara. F r o m 450 to 250 million years ago Africa was never very far from the South Pole. At that time violent earth movements caused the rocks of the Cape System to be strongly folded. Subsequent erosion has moulded the ranges which run parallel to the coast south of the Great Karoo. The Karoo gives its n a m e to a system of rocks which accumulated over m u c h of southern and central Africa during the Upper Carboniferous, Permian, Triassic and Lower Jurassic. The system includes glacial tillites, marine clays and coal-bearing continental deposits, debris eroded from the Hercynian fold mountains, deltaic and lacustrine sandstones, and enormous sheets of volcanic lavas. In Lesotho the lavas are as m u c h as 1800 m thick. In north Africa, the Continental Intercalaire, a system of rocks varying to a similar degree, but m u c h thinner than the Karoo, accumulated at a somewhat later stage. It includes the Nubian sandstone and other permeable water-bearing beds that underlie m u c h of the Sahara and are tapped by deep boreholes. During the late Mesozoic, Africa became separated from the rest of Gondwanaland. A s the oceans around Africa widened, freshwater and then marine deposits accumulated on the continental margins. Great fractures appeared in the east, letting d o w n blocks of Karoo sediments, which subsequently guided the evolution of the relief and drainage. Similarly in West Africa, the Benue fault trough opened from the Gulf of Guinea, penetrating far to the north-east into the Sanaran region. F r o m the Cretaceous (100 million years ago) onwards the geological history of the south-eastern part of the continent is quite different from that of the north-
west. The former was left high and dry, and only its margins were covered with marine sediments. The north and west were flooded by seas advancing south from Tethys in which thick layers of sandstone and limestone were laid down. During the late Cretaceous and Tertiary periods Africa moved north, slowly deforming the sediments in the Tethys ocean to produce the European Alps and related structures, including the Atlas. This Alpine deformation was associated with several faults with large horizontal movements. O n e such series of faults separates the folded rocks of the Maghreb from therigidSanaran block. South of the Atlas, the rock strata were not greatly contorted by the late Mesozoic and Tertiary earth movements, except in the Benue trough, where Cretaceous sandstones and clays were quite strongly folded and have since been dissected to give scarpland topography. Elsewhere, the basin form of certain depressions was accentuated, and stresses in the crust caused large-scale faulting. Evidence exists of successive uplifts of High Africa and of watershed areas throughout the continent during late Mesozoic and Tertiary times. The uplifted areas in High Africa form the Great African Plateau, which is the largest plateau in the world. Most of the plateau is more than 900 m above sea-level. Generally, the edge of the plateau forms the highest ground and is in the nature of a watershed between the headwaters of plateau rivers and those of the coastal drainage. In southern Africa the highest parts of the plateau rim are on the Drakensberg (Thabana-Ntlenyana, 3485 m , and m a n y points above 3190 m ) and the volcanic cone of M t Rungwe, 2961 m , in southern Tanzania. In between, the syenite mass of Mlanje in southern Malawi rises to 3000 m , the granite Namuli Peak in Mozambique to 2419 m and Inyangani in Zimbabwe to 2515 m . O n the western side of the subcontinent the highest points on the plateau are in Angola (Mt M o c o , 2620 m , Serra da Chela, 2300 m)^ and in the Auas highlands south of Windhoek (Molkteblick, 2485 m ) , but the highest point in the west lies outside the plateau where the Brandberg rises steeply from the N a m i b desert to an altitude of 2695 m . The original edge of the plateau probably at one time formed the coastal margin of the continent, but as streams cut back into the plateau edge and as the sea floor became exposed during uplifts of the land mass, the edge of the plateau no longer formed the coastline, but became a physical feature separating the plateau from the coastal region. In the course of time, with the continued retreat of the plateau edge, and with further exposures of the seafloor,the area between the plateau and the coast became in places so extensive that it can n o w no longer be considered coastal in character, but rather as a region marginal to the plateau, its inner areas removed from the coast in some places hundreds of kilometres.
Geology and physiography
In southern Africa the boundary between the plateau and areas marginal to it is generally called the 'Great Escarpment'. The latter is a variable feature, particularly in height, abruptness, and steepness of slope, depending chiefly on the configuration of the plateau itself, its rock formations, and on climate. In general it is abrupt, clifflike and linear where hard, resistant formations overlie soft ones. Where the rock is hom*ogeneous and easily decomposed the escarpment is an irregular feature with more gentle slopes. The Great Escarpment is most abrupt and prominent along the border of Natal and Lesotho, where it is capped by Stormberg lavas and is k n o w n as the Drakensberg. Elsewhere in southern Africa it is usually a well-defined feature, but in places it vanishes or is not very pronounced. Thus there is a 95 k m wide gap between the Koudeveld and Nieuweld M t s in Cape Province. North of the Swakop River in Namibia there is n o distinct edge to the plateau for 480 k m except for the western face of the Erongo Mts. In Angola north of the Huilla plateau the escarpment forms the watershed between the Cunene basin and the coastal drainage but does not form a very well-defined feature. North of the Natal Drakensberg the Great Escarpment is only intermittently conspicuous and n o plateau edge is discernible in the Limpopo valley, but in Z i m b a b w e the feature is again recognizable in the Melsetter-Chimanimani highlands and the Inyanga scarp. Further north, owing to advanced dissection and to complex trough faulting the position of the plateau edge is difficult to locate. T o the east of Lake Malawi it is possible that the N j o m b e highlands represent the position of the plateau edge in pre-rift valley times, but this is conjectural. In southern Africa two primary divisions of the great plateau can be recognized: the central or Kalahari basin, and the peripheral highlands, which are widest in the east and narrowest in the west. The Kalahari basin extends some 1930 k m from the Orange River to the southern Zaire watershed. Its greatest width is 1300 k m and it has an area of about 1640000 k m 2 . It is almost entirely covered with a mantle of sand, which extends across the watershed into the Zaire basin, and is probably the largest continuous surface of sand in the world. It is widely believed that each uplift of High Africa was followed by the cutting of an erosion surface graded to a lower base level than the one preceding it. S o m e geomorphologists have looked upon the continent's relief as consisting essentially of broad erosion levels and intervening erosional escarpments, of which the most striking of all is the Drakensberg in the Republic of South Africa. Dating of the erosion surfaces and the stages in the evolution of the relief is difficult, and the w a y in which the erosion surfaces have been cut is still uncertain. King (1967, 1978) groups the surfaces together into a few major 'cycles' and attributes them to pediplanation involving scarp retreat over long distances, but more
21
evidence is required before this interpretation of African landscape can be unreservedly accepted. The most striking feature of the relief of High Africa is therift-valleysystem, which extends from Turkey to Zimbabwe. In east Africa two fault systems can be distinguished. T h e eastern rift bisects Tanzania, cuts across the Kenya highlands to Lake Turkana (Rudolph), then turns north-east, splitting the Ethiopian plateaux and diverging into the m u c h wider trenches of the Afar depression, the R e d Sea and the Gulf of Aden. The western rift can be traced from the upper Nile and Lake Edward, through Lakes Kivu, Tanganyika and Malawi to the coast near Beira, with a branch extending along the Luangwa valley, the middle Zambezi and the southern margins of the Okavango swamps in Botswana. The general pattern of theriftvalleys seems to correspond with the lines of ancient structures in the crystallinefloor.In the southern part of the system, some of the troughs such as the Luangwa valley seem to have been subsequently eroded in relatively soft sediments that filled the troughs after they had been let d o w n between parallel faults at the end of Karoo times. They are essentially erosional features. In others, like the Malawi and Tanganyika troughs, the form of the valleys has been sharpened in late Tertiary and Quaternary times by dislocation along earlier fault planes, sometimes involving vertical movements of m a n y hundreds of metres. T h e separation of Arabia from Africa to form the R e d Sea and Gulf of A d e n has taken place within the last 15 million years, and locally, as in the Afar and Danakil depressions of Ethiopia, faulting has continued into present times. M a n y of the Indian Ocean and Atlantic islands are the fragments of old volcanoes that originated on the crests of mid-ocean ridges and were carried away from the crest as the sea floor continued to spread, but the Seychelles and Madagascar are isolated bits of continent. T h e oldest volcanic islands are the furthest from the ridge crests. A volcano on Réunion is still active and so are M o u n t Cameroun and a number of the volcanoes in the vicinity of the rift valley, notably the Virungas near Lake Kivu. The coastline of Africa, like that of Gondwanaland generally, is remarkably free from indentations. This is probably to be explained by its faulted character, by the lack of folding in late geological times, by continental uplift being dominant and by the deposition of riverborne sediments along the coast as sandbars and deltas during and since the rise in sea-level at the end of the last glaciation in high latitudes. Climatic conditions have varied all over Africa during the last million or more years of the Quaternary period. Nearly everywhere there is evidence that the climate has been both wetter and drier and warmer and colder in the past. There is, however, m u c h uncertainty concerning the precise pattern of change, even during the last 20000 years for which most evidence is available.
22
Environment, land use and conservation
The climatic sequence in the drier parts of north Africa subsequent to the melting of the ice-caps in Europe and North America about 15000 years ago appears to be fairly well documented. B y contrast, some conclusions about other parts of Africa, e.g. those of Livingstone (1967, 1975) on East Africa are not fully supported by the evidence cited, which is partly palaeobotanical (White, 1981). It is clear that the subject is m u c h more complicated than is commonly assumed, especially concerning the effects of climatic change on the distributions of animal and plant species, and hence on the floristic composition of plant communities. Although this subject is important in the understanding of African vegetation it lies outside the scope of the present work. Quaternary climatic fluctuation has also had profound indirect effects on vegetation through changes in soil, drainage systems and land-forms. Frequent mention is m a d e in the text of Quaternary aeolian sands and lacustrine clays. T h e influence of Quaternary climatic change on land-form has been described in some detail by Grove (1969) for the Kalahari region and Grove & Warren (1968) for the southern Sahara. Kassas (1953a, 1956a) has described the desert vegetation of Egypt and the Sudan in relation to land-form.
Although our understanding of African vegetation is greatly enhanced if it can be related to parent material and physiography, the influence of the former is usually indirect through its control of physiography rather than through the chemical nature of the soils to which it gives rise. There are, of course, exceptions. T h e extensive literature on heavy metal and other toxic soils has been summarized by Wild (1978). Such vegetation is comparatively localized and is only briefly mentioned in the present account. The distribution of saline soils is partly determined by geology in that they can occur in relatively wet regions around springs provided evaporation is sufficiently high, as in parts of the Mediterranean Region. In parts of east Africa, where the m e a n annual rainfall is 250-1000 m m , salts derived from volcanic deposits rich in sodium are deposited in lake basins andrivervalleys (see page 266). In the Zambezian region rocks containing the soda feldspar perthite locally giveriseto sodium soils, which, though not sufficiently saline to support halophytic communities, nevertheless carry distinctive vegetation (see page 94). The colonizing vegetation of recent lavaflowshas been described by Keay (1959a), J. Lebrun (1959, 1960¿), A . Léonard (1959) and Robyns (1932).
2
Climate and plant growth
Important general works include those by Aubreville (1949), S . P . Jackson (1962), B . W . Thompson (1965, 1966), Griffiths (éd., 1972), and I.J. Jackson (1977). The climate of the Zaire basin is summarized by Bernard (1945) and Bultot (1971-77), and of East Africa by Griffiths (1962). The rainfall m a p of the drier parts of East Africa and Arabia compiled by Griffiths & H e m m i n g (1963) is based partly on ecological information to compensate for the paucity of meteorological data. Swami (1973) has described moisture conditions in the savanna region of West Africa. Agroclimatological studies have been m a d e for the highlands of eastern Africa by Brown & Cochemé (1969), and for the drier parts of West Africa by Cochemé & Franquin (1967). Woodhead (1970) has used water balance in East Africa as a guide to site potential. Trochain (1952) has mapped the phytogeographic units of West Africa using bioclimatic data, and Papadakis (1966, 1970) has divided the world into climatic regions based on critical temperatures of certain cultivated plants and the water balance of the soil. This is the classification used in conjunction with the F A O - U n e s c o Soil Map of the World (1977). A more detailed survey of climate in relation to world vegetation has been edited byMonteith(1976). A m o n g more specialized studies, Trapnell & Griffiths (1960) have described rainfall in relation to altitude in Kenya. The importance of light rainfall in semi-arid areas is emphasized by Glover (P.E. Glover et al., 1962, J. Glover & G w y n n e , 1962). Rainfall variability is discussed by Nieuwolt (1972) for Zambia, and by Pennycuick & Norton-Griffiths (1976) for the Serengeti ecosystem, Tanzania. Tyson (1978) adduces evidence for a quasi-20-year oscillation in rainfall for the summer rainfall region for South Africa, and a 10-year oscillation for the all-seasons rainfall region of the southern Cape. T h e literature on mist precipitation and vegetation has been briefly reviewed by Kerfoot (1968). Walter (1936) has described the effects of fog on the vegetation of the N a m i b desert, and the mist oases of the eastern Sudan are the subject of publications by Troll (1935a) and Kassas (19566). Walter (1939; Walter & Walter, 1953) has also emphasized the importance of soil texture in relation to rainfall in the drier parts of southern Africa, and more generally (1955a), as has Smith (1949) for the Sudan. Included among the relatively few other works on water
relations of African plants are publications by Okali (1971) on some woody species of the Accra Plains, Ghana, by Ernst & Walker (1973) on the hydrature of trees in m i o m b o woodland, by Vieweg & Ziegler (1969) on the 'Resurrection Plant', Myrothamnus flabellifolius, and by Gaff (1977) on the poikilohydrous plants of southern Africa generally. The recent severe drought in the Sahel zone and the famine associated with it have stimulated several publications on desertification (e.g. Depierre & Gillet, 1971; Boudet, 1972; Delwaulle, 1973; Michon, 1973; W a d e , 1974). The general consensus is that destruction of vegetation by domestic animals and m a n has had far more effect than recent climatic deterioration. Boudet suggests that the brousse tigrée patterns of vegetation are probably brought about by wind action and sheet flow following degradation of the original vegetation. L . P . White (1971), however, believes that they are highly stable and have been in existence throughout the Quaternary period, but have migrated in response to successive climatic changes (but see page 27). The effect of wind on vegetation is described by Phot (1950ft, coastal vegetation in Senegal), Jenik (1968, tropical West Africa), Jenik & Hall (1966, effect of the harmattan in the Togo Mountains, Ghana), and Marloth (1907, South Africa). Tyson (1964) describes the berg winds of South Africa, which are usually characterized by a spectacular rise in temperature. Saboureau (1958) provides a graphic account of the destruction wrought by cyclones and floods on the vegetation of Madagascar, and Sauer (1962) deals with the influence of cyclones on coastal vegetation in Mauritius. Several workers have attempted to classify climate by means of indices based on selected factors which are assumed to be important in influencing plant growth, especially that of cultivated crops. The most widely used classifications of this kind are those of Koppen and of Thornthwaite. M o r e recently Holdridge has tried to define the world's major vegetation units in terms of logarithmic increases of temperature and rainfall units. The use in such systems of m e a n values which take no account of extremes was criticized by Moreau as long ago as 1938. The systems of Kóppen, Thornthwaite, and Holdridge are discussed in some detail for southern Africa by Schulze & M c G r e e (1978, q.v. for references). The application of Thornthwaite's system with various refinements to rangeland management in East Africa is
24
Environment, land use and conservation
described by Pratt & G w y n n e (1977). Emberger's climatic index is referred to in Chapter VII. A s Walter (1963) points out, it is not possible to express climate satisfactorily by figures or formulae, however complex, because of the seasonal rhythm of the most important factors, and variation from one year to another. B y contrast, diagrams, although far from perfect, can be used to summarize an enormous amount of relevant information, and permit rapid visual comparison of different stations and different vegetation types or different chorological and climatic regions. Walter (19556, 1959, 1963) has adapted a model first proposed by Gaussen (1955), and has published c. 10000 climatic diagrams in an atlas covering the whole world (Walter & Lieth, 1960-67; Walter, Harnickell & Mueller-Dombois, 1975). Walter's diagrams (see Fig. 3), besides summarizing up to eleven temperature parameters, show the annual march of m e a n monthly temperature and m e a n monthly rainfall, plotted on the same scale such that 20 m m of rainfall is equivalent to each increase of 10° C above zero. It has been empirically established that a relatively arid period prevails when the rainfall curve falls below the temperature curve, and that a relatively humid period occurs when the rainfall curve rises above the temperature curve. The vertical extent of the hatched and dotted fields in the diagrams provides an indication of the intensity of the humid or arid periods respectively. It must be emphasized, however, that these are relative values which apply only to the climatic type represented by the diagram. This is because it was necessary to indicate drought by the use of a temperature curve instead of an evaporation curve, since potential evaporation is measured only at very few stations, and measurements of radiation, wind, and air humidity are rarely available for estimating evaporation. The curves for temperature and potential evaporation often run parallel to each other, but these are not identical. In different climatic types the relation of temperature to potential evaporation is different and the difference increases with aridity. A drought period when shown on a climatic diagram is more severe the more arid the climate. This shortcoming of Walter's method is unavoidable where broad comparison on a continental scale is necessary, and is relatively unimportant if the objective is primarily to describe the climates of different vegetation types. The interpretation of plant growth, however, both indigenous and exotic, in terms of climate is another matter, and for this purpose all indices and diagrams are of limited value, and must be supplemented by direct physiological investigation. Unfavourable seasons caused by cold are also shown on Walter's diagrams. Months with a mean daily minim u m below 0 ° C are indicated by black blocks, and months with an absolute m i n i m u m below 0 ° C are hatched but frost m a y occur only in exceptionally cold years.
Additional climatic data are shown by numbers. In the tropics the monthly rainfall is often extremely high, and in order to simplify the diagram, the scale for rainfall above 100 m m per month is reduced 10 times. This area is shown in black and represents a perhumid season. ?
£
f1
?
! I ! I YANGÁMBI (487m) 244° 1964
34-9' 30-3'
30 C
LINDLEY (1524 m) 15-3° 647 , (13-48)
a b c d e f g h i j k I m n o q r
Station Altitude N u m b e r of years of observation (the first stands for temperature, the second for precipitation) M e a n annual temperature in ° C M e a n annual precipitation in m m M e a n daily m i n i m u m of coldest month Absolute m i n i m u m M e a n daily m a x i m u m of hottest month Absolute m a x i m u m M e a n daily range of temperature Monthly means of temperature (thin line) in ° C Monthly means of precipitation (thick line) in m m Arid period (dotted area) H u m i d period (hatched area) Perhumid area (black area), m e a n monthly rainfall over 1 0 0 m m (scale reduced ten times) Months with a m e a n daily m i n i m u m below 0 ° C Months with an absolute m i n i m u m below 0 ° C
F I G . 3. Climatic diagrams for Y a n g a m b i a n d Lindley
Climate and plant growth
In the equatorial region the daily variation in temperature is m u c h larger than the seasonal variation. Therefore additional figures are necessary: the m e a n daily m a x i m u m of the hottest month; the absolute maxim u m temperature; and the amplitude of the m e a n daily variation of the temperature. In order to facilitate comparison, the sequence of months for stations in the northern hemisphere starts with January, whereas for the southern hemisphere it begins with July. This means that the w a r m season is always in the middle of the diagram. Climatic extremes are often more important for plants than the means, and for agriculture and forestry, in particular, it is important to k n o w h o w often such extremes m a y occur. For this purpose climatograms are used. Their construction is nearly the same as that of ordinary climatic diagrams, except for the curves, which show the values for single years (preferably for a period of at least twenty years) instead of the means (see Walter, 1971, p. 56-57). Climatograms have been prepared for very few stations. Although they are indispensable for certain studies they are less essential for the purpose of broad regional comparison. Walter & Lieth have published more than 1000 climatic diagrams for Africa in their Weltatlas. They distinguish ten principal world climatic types, of which five occur in Africa, namely: I. Equatorial type, humid or with two rainy seasons. II. Tropical type with s u m m e r rain. III. Subtropical type, hot and arid. IV. Mediterranean type, with arid summer, and winter rains, frost rare. X . Mountain types. Each of these is subdivided and the boundaries of 51 subdivisions including transitional types are shown in the Weltatlas. O n m a p 3 of the publication of Walter, Harnickell and Mueller-Dombois (1975), 392 climatic diagrams for the African mainland and 14 for Madagascar and the Comores are superimposed in the approximate location of the meteorological station. T h e boundaries of the climatic types they represent are not shown on this m a p . The four main climatic regions in Africa and the transitions between them are, however, shown at a scale of 1:30000000 on their m a p 9.
25
A n enormous amount of climatic information is included in the Weltatlas and the Climate Diagram Map, and certain broad correlations between climate and vegetations are revealed by the latter. Nevertheless, it is still difficult to characterize the main phytochoria and vegetation types of Africa using these publications. For this reason, in the present work separate m a p s (Figs. 5-8 and 11-23) summarizing their climates have been prepared for all the major African phytochoria. For each phytochorion the climate is also briefly described. Elsewhere in the text climatic features of special significance are mentioned under individual vegetation types. Although it is relatively easy, using Walter's data, to characterize the climates of the major phytochoria, no attempt is m a d e here to produce a n e w bioclimatic synthesis, since it is apparent that the relationships between climate and vegetation are m u c h more complex than has generally been supposed. Thus, White (19786), w h o has m a p p e d the distributions of the Guineo-Congolian species of Diospyros, concluded that historic factors have played an important part. Also some features, such as dry-season atmospheric humidity, which do not contribute to the climatic diagrams, have a considerable influence on the vegetation. Similarly heat advection, as by the harmattan, is largely ignored by Walter's method. Relatively few attempts have been m a d e to relate growth and phenology to climate factors, other than crudely. A notable exception is the introductory text on tropical forest by L o n g m a n & Jenik (1974). T h e distribution of m i o m b o in relation to temperature and frost has been studied by Ernst (1971). Various aspects of phenology have been studied by Koriba (1958, tropics generally), Menault (1974, Ivory Coast), Yanney-Ewusie (1968, Ghana), M a d g e (1965, Nigeria), Njoku (1963, 1964, Nigeria), Huxley & van Eck (1974, Uganda), Kornas (1977, pteridophytes, Zambia) and Malaisse (1974, Malaisse et al, 1970, 1975, Zaire). The growth of two rapidly growing pioneer species of secondary forest, Trema orientalis (guineensis) (Coombe, 1960) and Musanga cecropioides (Coombe & Hadfield, 1962), has been analysed in great detail. In both species it was found that the rapid growth appears to be related to the prolonged and efficient development of new leafarea, rather than to a high rate of dry weight increase per unit leaf-area.
3
Soils
This section is intended to introduce botanists to literature they might find useful and to mention briefly some topics which recur at various places in Part Three, in the context of their associated vegetation. The soils of Africa have recently been mapped and classified twice for the whole continent. One system, the C C T A classification (D'Hoore, 1964), is local to Africa, created by scientists working in Africa, whereas the other (FAO-Unesco, 1974, 1977) tries to encompass the whole world. The C C T A system is summarized by D'Hoore (1968), A h n (1970, p. 213-19) and Young (1976, p. 236-40), and the F A O - U n e s c o system by Young (1976, p. 240-48). Information on the sou resources of tropical Africa is included in a symposium volume edited by Moss (1968). D'Hoore (1959) has briefly compared the soils of South America and Africa. Regional accounts include those by A h n (1970) on West Africa, Jones and Wild (1975) on West African savanna soils, and Scott (1962) on East Africa. M a n y references to individual countries can be found in the general works cited above but the following publications deserve special mention, in most cases because they attempt to relate vegetation to soils: A h n (1961, Ghana), Audry & Rossetti (1962, Mauritania), Ballantyne (1968, Zambia), Bawden & Carroll (1968, Lesotho), Bawden & Stobbs (1963, Botswana), Blair Rains & M c K a y (1968, Botswana), Brown & Young (1964, Malawi), Diniz (1973, Angola), H e m m i n g (1966, Somalia), H e m m i n g & Trapnell (1957, Kenya), Hopkins (1966, Nigeria), Latham & Dugerdil (1970, Ivory Coast), Milne (1947, Tanzania), Morison et al. (1948, Sudan), Perraud (1971, Ivory Coast), Pias (1970, Chad), Streel (1963, Zaire, Upper Shaba, Lufira), A . S . Thomas (1941, Uganda), Thompson (1965, Zimbabwe), Trapnell (1953, Zambia), Trapnell & Clothier (1937, Zambia), Trapnell et al. (1950, Zambia), Webster (in Chapman & White, 1970, Malawi), Wilson (1956, Zambia, Copperbelt) and Young & Brown (1962, Malawi). The soils of Zaire and R w a n d a and Burundi have been mapped in relation to vegetation in greater detail than those of other African countries. Twenty-six publications in the series Carte des sols et de la végétation du Congo belge et du Ruanda-Urundi (continued as Carte... du Congo, du Rwanda et du Burundi) deal with the following areas: — Zaire, Rwanda, Burundi (Sys, 1960, general soils map, 1:5000000)
— Rwanda, Burundi (Van Wambeke, 1963, soils) — Kaniama, Upper Lomami (Focan & Mullenders, 1955, soils, vegetation) — Mvuazi, Lower Zaire (Denisoff & Devred, 1954, soils, vegetation) — Ruzizi Valley (Germain et al., 1955, soils, vegetation) — Nioka, Ituri (Holowaychuk et al., 1954, soils, vegetation) — Mosso, Burundi (Bourbeau et al., 1955, soils, vegetation) — Yangambi: W e k o (Van W a m b e k e & Evrard, 1954, soils, vegetation) — Yangambi: Yangambi (Gilson et al, 1956, soils, vegetation) — Yangambi: Luanda (Gilson et al., 1957, soils, vegetation) — Yangambi: Y a m b a w (Van W a m b e k e & Liben, 1957, soils, vegetation) — Bugesera-Mayaga, Rwanda (Frankart & Liben, 1956, soils, vegetation) — Lufira Valley, Upper Shaba (Van W a m b e k e & Van Oosten, 1956, soils) — Lubumbashi, Upper Shaba (Sys & Schmitz, 1956, soils, vegetation) — Kwango (Devred et al., 1958, soils, vegetation) — Ubangi (Jongen et al., soils, vegetation) — Bengamisa (Van Wambeke, 1958, soils) — Lake Albert (Van Wambeke, 1959, soils) — Uele (Frankart, 1960, soils) — Kasai (Gilson & Liben, 1960, soils, vegetation) — 'Dorsale du Kivu' (Pécrot & A . Léonard, 1960, soils, vegetation) — Yanonge-Yatolema (Van Wambeke, 1960, soils) — Karuzi basin (Pahaut & Van der Ben, 1962, soils, vegetation) — Maniema (Jamagne, 1965, soils) — Tshuapa-Équateur (Jongen & Jamagne, 1966, soils) — Paysannat Babua (Frankart, 1967, soils) — Ubangi (Jongen, 1968, soils) — Upper Lulua (Gilson & Francois, 1969, soils) — Mahagi (Sys & Hubert, 1969, soils) — Lower Zaire (Compère, 1970, vegetation) — North Kivu and Lake Edward (Jongen et al., 1970, soils) M o s t of the m a p s are at scales of 1:50000 or 1:100000, but others range from 1:10000 to 1:1000000. All are in colour. General works on weathering and soil formation include those b y T h o m a s (1974) and N y e (1954/5). T h e origin of latérite is reviewed b y McFarlane (1976) and of duricrusts generally by Goudie (1973). S o m e aspects of
Soils
soil genesis in Central Africa are discussed by Ellis (1958), Webster (1960) and Paton (1961). W h e n soils and vegetation are mapped independently the results frequently do not correspond. There are several possible reasons for this. O n e is that the scale of variation of soil and vegetation is different. Another is that the vegetation is m u c h more attuned to the present-day climate than is the soil, some of which has its origin in a quite different climate from that n o w . Nevertheless, when soils and vegetation are studied jointly, especially in relation to the evolution of landscape and drainage patterns and the activities of soilforming animals, good, though sometimes unexpected, correlations are revealed. Studies along these lines are fewer than the purely descriptive and classificatory. Those mentioned below are of particular interest to the botanist. Trapnell (Trapnell, 1943; Trapnell & Clothier, 1938; Trapnell, Martin & Allan, 1950; see also Astle et al., 1969, and Webster, 1960) was one of the first to combine soil and vegetation in ecological survey. Although he published relatively little his influence has been considerable. Trapnell realized that, at least in Central Africa, climatic effects are often masked b y or subsidiary to those of geomorphology and age, and that the processes of intermittent continental uplift, peneplanation and faulting have greatly influenced soil formation and the differentiation of vegetation. H e used such topographic terms as 'Plateau', 'Upper Valley' and 'Lake Basin' to denote major soil groups in Zambia. Trapnell's units were based on the dominant, that is, the most extensive, vegetation-soil association in an area. They did not purport to describe fine detail nor to list all the minority vegetation-soil types within each mapping unit. Where such considerations are important the recognition of catenas and land-forms has proved valuable. The concept of the catena (Milne, 1935, 1936, 1947), which was originally defined 'as a certain sequence of soil profiles in association with a certain topography', is particularly valuable in explaining the relationships between soils and vegetation. Following Milne's example, several investigators in describing soil catenas have also included vegetation. O f special interest to the botanist are the catenas described by Bourguignon et al. (1960), Duvigneaud (1953), Lawson et al. (1968, 1970, Ghana), Morison et al. (1948, Sudan Republic), Radwanski & Oilier (1959, Uganda), Watson (1964, 1965, Zimbabwe), Webster (1965, Zambia) and Williams (1968, stabilized sand dunes, Gezira, Sudan Republic). A repeating pattern of vegetation change analogous to that associated with soil catenas occurs at m a n y localities in arid and semi-arid regions. Vegetation stripes running parallel to the contours occur on virtually flat to gently sloping surfaces. They are very conspicuous on aerial photographs, and have variously been described as vegetation arcs, bands orripplesand as 'brousse tigrée' (Boaler & Hodge, 1964; Clos-Arceduc, 1956; H e m m i n g , 1965; Macfadyen, 1950; L . P . White,
27
1970, 1971; Wickens & Collier, 1971; Worrall, 1959). The vegetation of the bands is denser, taller and physiognomically more complex than that of the intervening lanes, which are sometimes virtually devoid of vegetation. According to L. P. White, in most of the examples described, the stripes are not due to gross soil differences. W h a t differences there are between the soils of the two phases can be accounted for by the influence of the vegetation itself. For certain purposes of land classification for management planning the catena is too restrictive, and in recent years several studies, inspired by the work of Bourne (1931), have used a framework of land classes based on physiography, within which the different soil and vegetation units are subordinated. This approach has been widely used in Australia. In Africa it provides the bases of the land-resource studies of the British Ministry of Overseas Development. For a general discussion see Astle et al. (1969). This method recognizes that in any one landscape there are only a few kinds of terrain, each with its particular combination of land-forms, rocks, soil and vegetation. These few terrain types recur in association with one another in the landscape to give a more or less regular pattern always with the same interrelations. A new landscape is recognized where there is a change either in the terrain types or in the relations between them. This approach, which is of wider application than that of the catena, is more appropriate to rapid survey of extensive little-known areas. Astle et al. found that the patterns of landscape are easily distinguished on air photographs and their component parts consistently recognizable under the stereoscope. They recognize composite landscape patterns called Land Systems, the component parts of which are Land Facets. In their survey of the Luangwa Valley, Astle et al. describe the soils and vegetation of forty-six Land Facets which are grouped in nine Land Systems. Similar methods have been used in Botswana (Bawden & Stobbs, 1963; Bawden, 1965), western Kenya (Scott et al, 1971), Lesotho (Bawden & Carroll, 1968), Nigeria (Bawden & Tuley, 1966), Swaziland (Murdoch et al, 1972), and U g a n d a (Oilier et al, 1969). The part played by termites in tropical soil formation is certainly considerable but its extent is still controversial. Termites dominate the macrofauna of tropical and subtropical soils in the w a y that earthw o r m s dominate temperate soils, but their influence descends to considerably greater depths. Termites transport large volumes of mineral and organic matter, both vertically and laterally, and in m a n y places the top metre or so of soil has been radically modified by termite action. T h e nests of some species of termite are contained in m o u n d s of earth built up above the ground surface. The largest mounds, which m a y be up to 9 m in height, represent the work of species of Macrotermes. They are particularly conspicuous on the unrejuvenated plateau surface in the heart of the Zambezian Region, and most of the recent literature on the soil-forming
28
Environment, land use and conservation
activities of termites refers to them (Hesse, 1955; Meikeljohn, 1965; Sys, 1955; Watson, 1962a, 1967, 1969, 1974a, 19746). The soil within a termite m o u n d is very different from that surrounding it. Usually there is more clay and less coarse sand. The p H is nearly always higher and the content of carbon, nitrogen and exchangeable bases, especially calcium, is greater. T h e microflora is also distinctive. There are more cellulose decomposers, denitrifiers, ammonifiers and nitrifiers, but fewer nitrogenfixing bacteria of the genera Beijerinckia and Clostridium. Termite mounds are subjected to less leaching than the surrounding soils. This should favour the retention of bases within the mounds, but it does not explain h o w they enter. T h e work of Trapnell et al. (1976) indicates that bases removed from the soil by the woodland canopy become concentrated in termitaria by the termites themselves which feed largely on dead wood and Utter. T h e vegetation of large termite mounds, which is very different from that of the surrounding soil, is briefly described at appropriate places in Part Three. The soils of the Zambezian Region are more diversified than those of some other parts of Africa and their influence on vegetation has been studied in greater detail. T h e extensive literature on the vegetation of heavy metal and other toxic soils, which in Africa have been chiefly studied in the Zambezian Region, has been reviewed by Wild (1978). The distinctive soils of mopane woodland and other types of Zambezian vegetation are discussed in Chapter II. Savory (1963) has shown that the distribution and height of the dominants of m i o m b o woodland in Zambia are closely correlated with effective soil depth. Elsewhere, root systems have been studied by Huttel (1969, 1975, Ivory Coast), Okali et al. (1973, Ghana) and Glover (1950-51, Somalia). Kerfoot (1963) has published a short review.
Experiments (Grant, n.d.) have shown that the very acid infertile Kalahari sands of Zimbabwe are deficient in boron and sulphur. S o m e soils in the Cape Region are also deficient in trace elements and even indigenous species show symptoms of nutritional imbalance (Schütte, 1960). There is evidence that sclerophylly is sometimes associated with nutrient deficiency, especially of phosphorus (Loveless, 1961, 1962; Beadle, 1966, 1968; Grubb & Tanner, 1976; Grubb, 1977), but little information is available from Africa. The effects on soil of shifting cultivation have been described by N y e & Greenland (1960) and Vine (1968). The former publication remains the most important work in English on soil fertility in the tropics. Nitrogen supply in the forests and savannas of West Africa has been studied by de R h a m (1974). Milne & Calton (1944) discuss the effect of clearing vegetation on soil salinity in a semi-arid part of Tanzania. T h e influence of annual burning on soil structure and fertility is dealt with by M o o r e (1960, derived savanna zone, Nigeria) and Trapnell et al. (1976, Zambia). Soil factors affecting the distribution of vegetation types and their utilization by wild m a m m a l s in East Africa are described by Anderson & Talbot (1965, Serengeti Plains) and Anderson & Herlocker (1973, Ngorongoro crater). S o m e leguminous trees are believed to increase soil fertility. In the Sudan, according to Radwanski and Wickens (1967), the yields of sorghum and other crops planted under Acacia albida are considerably greater than elsewhere. T h e pod and leaf-fall and perhaps the dung and urine of cattle which eat the pods and seek the shade of the trees increase the supply of nutrients and improve the physical condition of the soil. Dancette & Poulain (1968) describe a similar study of Acacia albida in Senegal.
4
Animals
A s explained in the general Introduction, the aim of this book is primarily descriptive, namely to provide a classificatory framework within which more detailed and localized studies, both of plants and animals, can be conducted and compared. The classification, however, is based on plants alone. In this context animals have been deliberately left out. This is because their habitats are too imperfectly correlated with vegetation types to provide useful diagnostic features. Their ranges either greatly transgress the boundaries of vegetation types or, alternatively, w h e n they are confined to an individual type, they usually occupy only part of it. Theoretical concepts such as the biocoenose, which attach equal importance to animals and plants, are useful, indeed essential, for the understanding of vegetation, but they are of little classificatory value. The interpretation of vegetation is another matter. Until about twenty years ago, the importance of the interaction of plants and animals in shaping African vegetation had, with few exceptions, been gravely neglected by botanists and zoologists alike. Recently, however, there has been m u c h activity in this field, especially by zoologists, but progress has been patchy, with most effort concentrated on large m a m m a l s , chiefly in East Africa. Detailed consideration of the effects of animals on vegetation lies outside the terms of reference of this book, even if sufficient information for a general synthesis were available, which is not the case. Nevertheless, where animals greatly influence vegetation this is briefly mentioned in the text. In an attempt to compensate for the omission just mentioned, this chapter on 'Animals' is mainly intended to introduce botanists to zoological literature of botanical interest. It is meant to be no more than an entrée to the subject, not a complete review. General works are few, but Cloudsley-Thompson (1969) and O w e n (1976) have respectively written introductions to the zoology and animal ecology of tropical Africa. Curry-Lindahl (1968) deals with zoological aspects of the conservation of vegetation in tropical Africa. Petersen & Casebeer (1971) have compiled a bibliography relating to the ecology of East African large m a m m a l s . Most other general works are devoted to single taxonomic groups of animals, especially m a m m a l s , more rarely birds. The books by Delany & Happold (1978) on
m a m m a l s , b y Leuthold (1977) on ungulates, and b y Moreau (1966) on the birds of Africa include m u c h of interest to the botanist, as does Kingdon's (1971-77) encyclopaedic work on East African m a m m a l s . Bigalke's treatment (1978) of the biogeography and ecology of the m a m m a l s of southern Africa is particularly valuable for its references to the literature, as is the review b y Bourlière & Hadley (1970) of the ecology of tropical savannas. O n a more local scale, Rosevear's (1953) check-list provides m u c h information on the distribution of Nigerian m a m m a l s in relation to vegetation. Bourlière & Verschuren (1960) have published a monograph on the ecology of ungulates in the Albert National Park. Most of the publications mentioned below deal with a single animal species or a few related species. Relatively few deal with animal communities in relation to plant communities. Important works a m o n g the latter are those by: — Chapín (1932,1939) on the birds of Zaire. — Moreau (1935a) on the birds of the Usambara Mts, Tanzania. — Fraser Darling (1960) on the ecology of the Mara Plains in Kenya. — Lamprey (1963, 1964) on ecological separation and population dynamics of the large mammal species in the Tarangire G a m e Reserve, Tanzania. — Coe (1967) on the vertebrate fauna of the Afroalpine zone in Kenya. — Anderson & Herlocker (1973) on soil factors which affect the distribution of vegetation types and the animals which use them in Ngorongoro Crater, Tanzania. — Sinclair & Norton-Griffiths (1979) on the dynamics of the Serengeti ecosystem in Tanzania. Aco*cks (1979) has attempted a reconstruction of the vegetation of the whole of the drier half of South Africa in relation to its indigenous fauna, as it was before the advent of the European. In recent years m u c h work has been done on the larger m a m m a l s , chiefly in relation to their feeding behaviour, population dynamics and influence o n vegetation in national parks and g a m e reserves. T h e pioneer publications of Eggeling (1939), Mitchell (1961a), Walter (1961) and Cornet d'Elzius (1964), which emphasized the potentially far-reaching effects of g a m e animals on vegetation, have been followed b y numerous detailed studies of several animal species, and
30
Environment, land use and conservation
by monographie treatments of the elephant (Wing & Buss, 1970; L a w s et al., 1975) and the buffalo (Sinclair, 1977). M a n y publications deal with the feeding ecology of herbivores, for example:
— Agnew (1968, Tsavo National Park (East), Kenya). — Buechner & Dawkins (1961, elephant, Murchison Falls National Park, Uganda). — Douglas-Hamilton (1973, elephant, Lake Manyara, Tanzania). — Glover (1963, elephant, Tsavo). — Glover & Wateridge (1968, cattle and wild ungulates as a cause of terrace erosion). — Harrington & Ross (1974, elephant, Kidepo Valley National Park, Uganda). — Kortlandt (1976, elephant, Tsavo). — Lamprey et al. (1967, elephant, Serengeti National Park, Tanzania). — Laws (1970a, 19706, elephant, East Africa). — Penzhorn et al. (1974, elephant, Addo Elephant National Park, Eastern Cape Province, South Africa). — Thomson (1975, elephant in Brachystegia boehmii woodland, Chizarira G a m e Reserve, Zimbabwe). — Van W y k & Fairall (1969, elephant, Kruger National Park, Transvaal). — Watson & Bell (1969, elephant, Serengeti).
— baboon (Lock, 19726). — black rhinoceros (Goddard, 1968,1970). — buffalo (Vesey-FitzGerald, 1969, 1974a; Leuthold, 1972; Sinclair & Gwynne, 1972; Grimsdell & Field, 1976). — buffalo, hippopotamus, Uganda kob, topi, warthog, and waterbuck (Field, 1972). — Chanler's mountain reedbuck (Irby, 1977). — duiker (Wilson & Clarke, 1962; Wilson, 1966). — elephant (Napier Bax & Sheldrick, 1963; Field & Ross, 1976). — fringe-eared oryx (Root, 1972). — gerenuk (Leuthold, 1970). — giraffe (Innis, 1958; Foster, 1966; Foster & Dagg, 1972; Leuthold & Leuthold, 1972; Field & Ross, 1976). — greater kudu (Wilson, 1965). — hares (Lepus capensis, L. crawshayi; Pronolagus crassiIn an attempt to understand the long-term implications caudatus, Stewart, 1971o-c). of the recent impact of large m a m m a l s o n the vegetation — hippopotamus (Field, 1970; Lock, 1972a). of reserves, several studies of the m o v e m e n t , habitat — impala (Stewart, 1971d; Rodgers, 1976). utilization, biomass, density, mortality, age structure — lechwe (Vesey-FitzGerald, 19656). and population dynamics, again especially of the eleph— lesser kudu (Leuthold, 1971). ant, have been m a d e , including those by: — primates generally (Clutton-Brock, 1977, éd.). — rock hyrax (Sale, 1965). — Bourhère (1965, ungulates generally). — rock hyrax and tree hyrax (Turner & Watson, 1965). — Coe et al. (1976, large African herbivores). — sitatunga(R. Owen, 1970). — Corfield (1973, elephant). — waterbuck (Kiley, 1966). — Lamprey (1964, large mammals generally). — wildebeest (Talbot & Talbot, 1963). — Leuthold (1976, elephant). — wildebeest and zebra (Owaga, 1975). — Leuthold & Leuthold (1976, ungulates generally). — wildebeest, zebra, and hartebeest (Casebeer & Koss, — Leuthold*Sale(1973, elephant). 1970). — wildebeest, Thomson's gazelle, Grant's gazelle, topi, and — Olivier & Laurie (1974, hippopotamus). impala (Talbot & Talbot, 1962). — Sinclair (1974, buffalo). — various ungulates (Pienaar, 1963; Gwynne & Bell, 1968; — Western & Sindiyo (1972, black rhinoceros). Stewart & Stewart, 1971; Pratt & Gwynne, 1977). N o t all degradation in reserves is brought about by large m a m m a l s , and in some cases their influence is indirect. T h e grazing succession throughout the year of the eight That fire is often involved in the destruction of c o m m o n e s t large herbivores in the R u k w a Valley, vegetation cannot be denied, though its influence varies Tanzania, namely elephant, buffalo, hippopotamus, locally, and interactions between fire and elephants puku, topi, zebra, bohor reedbuck and eland, has been deserve further study. According to Verdcourt (in litt. 18 described by Vesey-FitzGerald (1960, 1965a), w h o has D e c e m b e r 1978) elephants in K e n y a have sometimes also (19736, 1973c) studied browse production and been blamed for the extensive depredations of charcoalutilization in Tarangire and Lake M a n y a r a National burners. Parks. His surveys indicated that at the time of the study the animals, principally elephant, rhinoceros and giraffe, E v e n in areas such as the Akagera National Park in were utilizing only half of the browse available. R w a n d a , where elephants are virtually u n k n o w n today, a high proportion of trees are felled by wind or damaged W h e r e large m a m m a l s , especially the elephant, have by lightning, and so simulate elephant d a m a g e (Spinage been protected in recent years, they have often shown a & Guiness, 1971). dramatic increase in numbers, sometimes accompanied by wholesale destruction of vegetation with concomitant In areas of low rainfall (c. 350-400 m m per year), landscape change (see page 114). T h e extent to which especially in basins of closed drainage, elephants might such population changes can be regarded as 'natural' play a catalytic rather than a primary role in the decline and the desirability of exerting artificial control are of w o o d y vegetation (Western & V a n Praet, 1973). Thus, still matters of controversy and have given rise to an for the Amboseli G a m e Reserve in K e n y a , Western extensive literature. & V a n Praet cite evidence which indicates that cyclical climatic change is responsible for a cyclical alternation Publications which discuss large m a m m a l s as agents of Acacia xanthophloea woodland and treeless haloof habitat and landscape change include those by:
Animals
phytic communities dominated by Suaeda monoica. D u r ing the wet phase of the cycle the water-tablerisesby as m u c h as 3.5 m , and the capillary fringe introduces a high level of soluble salts into the rooting horizon of the Acacia xanthophloea trees, which ultimately die and fail to regenerate. Elephants merely hasten death brought o n by other causes. Notwithstanding the fact that trees are destroyed by fire, charcoal-burners, wind, lightning and changes in soil salinity, m u c h degradation must be attributed to elephants and other large m a m m a l s . It w a s formerly assumed that elephant/ forest systems possess a stable equilibrium point, and that rapid decline of vegetation only occurs w h e n this equilibrium is displaced by m a n , leading to local high densities of elephants. Caughley (1976) has proposed an alternative hypothesis. H e believes that the relationship is a stable limit cycle in which elephants increase while thinning the forest, and then decline until they reach a density low enough to allow resurgence of the forest. His hypothesis is incomplete since he does not mention the possible effects of cyclical climatic change o n the elephant/ vegetation cycle nor does he consider the relative importance of decline in elephant numbers due to natural catastrophic mortality and migration. H e does, however, discuss modifications brought about by m a n . Caughley's evidence comes from the Zambezi Valley in Zambia, where the uneven distribution of age classes of elephantdamaged baobab (Adansonia digitata), and m o p a n e (Colophospermum mopané) trees, suggests a cycle of the order of 200 years. A cyclical relationship between animal numbers and vegetation, but in this case controlled by rainfall, has also been postulated by Phillipson (1975), w h o related elephant mortality in Tsavo National Park (East) in K e n y a to estimated primary production. H e concluded that 'carrying capacity' declines markedly approximately once every 10 years for large m a m m a l s other than elephant, and only once every 43-50 years is there a change sufficiently marked to result in a crash in elephant numbers. There is n o single level of population at which a steady state is possible. Nearly all the literature o n elephant ecology refers to East and southern Africa. Little has been published o n the Guineo-Congolian Region, but elephants are k n o w n to retard succession of vegetation at places where they bathe and drink. In Zaire, for instance, the vegetation near elephant baths dominated by Rhynchospora corymbosa might represent an elephant-maintained subcümax (Léonard, 1951). T h e possibility of domesticating wild m a m m a l s for meat production is discussed b y D a s m a n n (1964) and Parker & G r a h a m (1971). Despite promising beginnings g a m e ranching in most places has failed as an economically viable agricultural practice. Alternative ways of cropping wild herbivores are considered by Pratt
31
& G w y n n e (1977) for East Africa and b y Huntley (1978) for southern Africa. Notwithstanding the considerable importance of fruit and seed dispersal by birds and m a m m a l s the subject has been little studied. T h e few publications include those by: — Burtt (1929, 28 plant species by 9 m a m m a l and 4 bird species). — Clutton-Brock (éd., 1977, primates). — Gwynne (1969,/4cac/a by ungulates). — Hladik & Hladik (1967, primates, Gabon). — Jenik & Hall (1969, Detarium microcarpum by elephants). — Kingdon (1971-77, mammals generally). — Lamprey (1967, Acacia by ungulates; Commiphora by birds, baboons, and monkeys). — Lamprey et al. (1974, Acacia tortHis by elephant, impala, dikdik, and Thomson's gazelle). — Leistner (19616, Acacia erioloba by elephant, giraffe, black rhinoceros, gemsbok, and eland). — Phillips (19266, forest trees in the Knysna region, Cape Province, South Africa by wild pig). — Van derPijl (1957, bats generally). — Wilson & Clarke (1962, Pseudolachnostylis maprouneifolia by the c o m m o n duiker). T h e effects o n vegetation brought about by s o m e groups of insect are n o less important than those of m a m m a l s , both directly (locusts; a r m y w o r m , E d r o m a , 1977) and indirectly through soil formation (termites), or through population control of m a m m a l s including m a n (tsetse fly). For each group there is an extensive specialized literature. Only a few of the m o r e general works can be mentioned here. T h e habitats of the Desert Locust are described by Guichard (1955) and by H e m m i n g & S y m m o n s (1969) and those of the R e d Locust by Backlund (1956), VeseyFitzGerald (1955a, 1964) and Rainey et al. (1957). T h e interrelationships of vegetation and termites have been described by Murray (1938) for South Africa, by Wild (1952a, 1975) for Z i m b a b w e , by Fries (1921) and Fanshawe (1968) for Zambia, and by Malaisse (1976a) for Zaire (Upper Shaba). T h e last author (Malaisse, 1978¿>) has also published a well-documented review of the termite m o u n d as an ecosystem for the whole of southern Africa. References to the soil-forming activity of termites are given in Chapter 3. Important publications o n the tsetse fly include those of Goodier (1968), N a s h (1969), Ford (1971) and Ormerod(1976). Despite a recent renewed interest in symbiotic relationships between ants and plants few publications deal with Africa. T h e association between ants and gall Acacias has been studied b y B r o w n (1960), M o n o d & Schmitt (1968), Hocking (1970, 1975), and Foster & D a g g (1972). T h e protection of the rain-forest tree Barteria fistulosa b y Pachy sima ants is described by Janzen (1972).
5
Fire, land use and conservation
Fire The effects of fire on vegetation axe often referred to in publications mainly concerned with other matters. Those mentioned below deal exclusively or principally with the subject, mostly with fires started deliberately or accidentally by m a n , but Komarek (1964, 1972) reviews the occurrence of fires caused by hghtning. It is generally agreed that frequent uncontrolled fires are harmful both to vegetation and soil, and that for some purposes controlled burning is beneficial. Some controversy, however, still surrounds the precise regime to be followed, and the long-term effects are often uncertain. In this account no attempt is made to resolve these problems. The principal literature is merely referred to. Other references and information are given in Part Three in relation to individual vegetation types. General works dealing with the ecological effects of fire and its use in land management include those by Humbert (1938), Bartlett (1956), Ahlgren (1960), West (1965), Daubenmire (1968), and Glover (1968, 1972), and for Africa only, Guilloteau (1957) and Phillips (1965, 1968, 1972, 1974). West Africa is covered by Scaëtta (1941), Viguier (1946), Pitot (1953) and Rose Innes (1972), southern Central and eastern Africa by V a n Rensburg (1972), and the Mediterranean Region by Naveh(1974). The role of fire in individual territories has been described by Lamotte (19756) and Monnier (1968) for the Ivory Coast, by Hopkins (1963, 1965, 1978c, 1979; White & Werger, 1978), based in part on a large number of detailed published and unpublished distribution m a p s of individual species, has confirmed the general validity of the phytochoria, though, not unexpectedly, a few minor emendations have been indicated. It should be emphasized that the distributional data which confirm the boundaries on the chorological m a p were not used in the compilation of the vegetation m a p . Most, in fact, were not then available. This concordance of chorology and physiognomy is doubly significant. It both helps to confirm the objectivity of the mapping units and provides an objective means of designating and differentiating vegetation types of broadly similar physiognomy which occur in differentfloristicregions. It also can provide a 'shorthand' reference to the entire physiognomy of a vegetation type, since the latter is determined by its entire flora. Reference to sclerophyllous vegetation should m a k e this clear. Since the time of Schimper (1898, 1903) the vegetation of the various widely separated regions with a Mediterranean climate has been regarded as equivalent both in physiognomy and ecology: 'All districts agreeing
41
with the Mediterranean coast as regards the distribution in time of the rainy and dry seasons repeat in their vegetation essential oecological features of the Mediterranean vegetation.' These regions 'are the h o m e of evergreen xerophilous plants which, owing to the stiffness of their thick leathery leaves, m a y be termed sclerophyllous woody plants' (Schimper, 1903). T h e term sclerophyll has been subsequently extended to similar vegetation in non-Mediterranean climates, such as parts of Australia (see Seddon, 1974) and on high tropical mountains. The leaves of some plants on the latter, however, have some distinctive anatomical features and Grubb (1977) refers to them as 'pachyphylls'. It would be foolish to deny the striking similarities in leaf-form in the different regions with a Mediterranean climate, but n o less so to ignore the important differences. C o d y & M o o n e y (1978), w h o have m a d e an interesting though limited comparison of the Mediterranean ecosystems of the five main areas in which they occur, found that the structure of the communities, the successional relationships and the growth rhythms all differ substantially a m o n g regions, with South Africa being the most dissimilar. In the Cape Region there is more summer rainfall than elsewhere and the main growth period is not confined to the spring but continues through the summer. The Cape vegetation has a greater diversity of shrubby forms, and is rich in bulbous plants, but poor in annuals and vines. A more detailed comparison would doubtless reveal other differences. Furthermore the successional status of 'Mediterranean' shrubland is different in the different areas. Thus, in the Cape Region 'fynbos' (Chapter V ) nearly everywhere represents the climax. True forest, other than some small areas of scrub forest, only occurs as enclaves where s u m m e r rainfall or cloud profoundly modifies the Mediterranean climate; floristically it is of Afromontane affinity. B y contrast, in the Mediterranean basin evergreen forest dominated by Mediterranean endemic species is the regional climax. The ecological significance of sclerophylly has long been and still remains a subject of controversy (for recent reviews see Seddon, 1974; Grubb, 1977; Cody & M o o n e y , 1978). There can be little doubt, however, that in different places different combinations of environmental factors are responsible. F r o m the above it is clear that sclerophyllous shrubland in the different regions shows appreciable differences in physiognomy, related in part to different environmental conditions which are still imperfectly understood. This poses considerable problems in classification. O n the one hand, it is clearly inappropriate that vegetation as different as that of the Mediterranean and Cape Regions should go under the same n a m e , e.g. the 'Hard-leaved scrub bushes (macchia)' of Schmithiisen (in F A O - U n e s c o , 1977). O n the other hand, in view of our incomplete knowledge of the physiognomic differences and particularly of their adaptive role, the selection of discriminating epithets is difficult. However,
Regional framework, classification and mapping units
42
if the sclerophyllous formations in the different floristic regions are designated and distinguished by the names of those regions, the arbitrary selection of imperfectly understood characters can be avoided. This applies equally to all other multiregional formations. Hence, the n a m e of the phytochorion can serve to e m b o d y and provide a key to all the diagnostic features of its vegetation types. In the main text (Part Three) the descriptions of individual vegetation types include as m u c h physiognomic information as was readily available, but published data are often sparse. Sometimes I have had to rely almost exclusively on m y o w n field records. A n additional advantage of grouping the main vegetation types under the regional phytochoria is that it permits a more effective treatment of mosaics, continua and transitions, and also of the dynamic relationships of important regional types which have suffered degradation at the hands of m a n . This can be exemplified by the Sudanian and Zambezian Regions, which have broadly similar climates. In both regions, woodland, broadly comparable in physiognomy andfloristiccomposition, is the most widespread type of vegetation. T h e Sudanian Region, however, belongs to ' L o w Africa' (Chapter 1), whereas the Zambezian Region belongs to 'High Africa' and hence has a more diversified physiography and climate. This is reflected in a wider range of vegetation types. In the Zambezian Region distinctive types of dry evergreen forest, dry deciduous forest, thicket and edaphic grassland are m u c h more widespread than their rather different analogues in the Sudanian Region, and this is only partly due to the effects of m a n . In the Zambezian Region the subordinate types form c o m plicated mosaics with the regional woodlands and have complex dynamic relationships with them. T h e pattern
itself is more important than the sum of its parts and is best described in a regional context.
The main phytochoria The chorological m a p mentioned above differs from earlier m a p s in two main respects. First, the phytochoria are based on richness (or otherwise) of their endemic floras at the species level. Second, no attempt is m a d e to carve up Africa into mutually exclusive areas which are themselves subdivided in a hierarchical manner (Regions, Sectors, Domains, Districts, etc.). Different parts of Africa differ greatly in theirrichnessin endemic species and in the distribution patterns shown by the latter. For this reason a flexible, non-hierarchical system is proposed (for a fuller account see White, 1979). It recognizes at the rank of Region four fundamentally different types of phytochorion, the salient features of which are embodied in the terminology employed (Table 2). A s provisionally defined (White, 1979), a Regional Centre of Endemism is a phytochorion which has both more than 50 per cent of its species confined to it and a total of more than 1000 endemic species. All the phytochoria designated above as Regional Centres of Endemism appear to fulfil these criteria with the exception of the Sudanian Region, the status of which is still uncertain. T h e Regional Centres of Endemism are separated by transition zones. If the latter are comparable in size to the former they should be named and given comparable rank. T h e transition zones between the Cape and K a r o o - N a m i b Regions and between most islands of the Afromontane Region and their surrounding lowland phytochoria are too narrow to justify this. B y contrast
T A B L E 2. The main phytochoria of Africa and Madagascar A Africa 1 REGIONAL CENTRES OF ENDEMISM:
I Guineo-Congolian II Zambezian III Sudanian IV Somalia-Masai V Cape VI Karoo-Namib VII Mediterranean
B Madagascar 2 ARCfflPELAGOLIKE CENTRE OF ENDEMISM:
VIII Afromontane
3 ARCHIPELAGOLIKE CENTRE OF EXTREME FLORISTIC IMPOVERISHMENT:
IX Afroalpine
4 REGIONAL TRANSITION ZONES AND MOSAICS:
X
Guinea-Congolial Zambezia XI Guinea-Congolial Sudania XII Lake Victoria XIII ZanzibarInhambane XIV Kahalari-Highveld XV TongalandPondoland X V I Sahel XVII Sahara XVII Mediterranean/ Sahara
1 REGIONAL CENTRES OF ENDEMISM:
XIX East Malagasy X X West Malagasy
Regional framework
the Guinea-Congolia/Sudania, Guinea-Congolia/Zambezia and the Kalahari-Highveld transition zones are larger than some Regional Centres of Endemism and need to be named. These three transition zones have few endemic species and the majority of their species also occur in adjacent phytochoria. T h e transition from the Mediterranean Region to the tropicalflorais more complex, and can conveniently be divided into three Regional transition zones, the Sahara, the Sahel and the Mediterranean/ Sahara transition zones. The last two have impoverished floras and few endemic species. Endemism is appreciably higher in the Sahara, but both the total and percentage of endemic species are too low for the Sahara to qualify as a Regional Centre of Endemism. Furthermore the Sahara forms a transition between two great floristic kingdoms, the Holarctic and the Palaeotropic, whose generic floras, subcosmopolites apart, are almost totally different.
43
T h e transition zones mentioned above show a simple gradual replacement of one flora by another which is little complicated by endemism. The three Regional Mosaics, however, are transitional in a more complex manner. In all three the vegetation forms a mosaic of different physiognomic types with different floristic relationships. The Lake Victoria Regional Mosaic has few endemic species, whereas endemism in the Zanzibar-Inhambane and Tongaland-Pondoland Regions is relatively high. T h e vegetation of the regional phytochoria is described in Part Three, where the mainfloristicfeatures of each phytochorion are briefly summarized. The figures relating to the size of the flora and the degree of endemism are estimates based on available information, m u c h of it unpublished (for discussion see White, 1979). Further work will certainly modify the details considerably, though the overall pattern seems to be well established.
7
Classification
Introduction
Introduction
Description of the main vegetation types Forest Woodland Bushland and thicket Shrubland Grassland Wooded grassland Desert Afroalpine vegetation Scrub forest Transition woodland Scrub woodland Mangrove Herbaceous fresh-water swamp and aquatic vegetation Saline and brackish swamp Bamboo Anthropic landscapes
The broad principles underlying the present classification are explained in the Introduction and in Chapter 6. Because these principles are somewhat different from those of most other classifications it was originally intended to include a longer discussion on classification in general. But this was precluded for lack of space. Nevertheless it might be useful to mention in this context the following publications, most of which are not discussed elsewhere: Aubréville (1951), B a m p s (1975), Beard (1978), Cain (1950), Chevalier (1953a, 19536), Dansereau (1951), D a s m a n n (1972, 1973a, 19736), D r u d e (1913), D u Rietz (1931), Emberger, Mangenot & Miège (1950a, 19506), Gaussen (1955, 1958), Kinloch (1939), Kiichler (1947, 1949, 1950, 1960, 1967, 1973), Plaisance (1959), Poore (1962, 1963), Redinha(1961), Robyns (1942), Schnell (1970-71,1977), Udvardy (1975, 1976), Walter (19766), Walter & B o x (1976) and W e b b (1954). In this chapter the classification is merely briefly considered in relation to earlier classifications of African vegetation. T h e distinguishing features of the sixteen major formations are also described, and an outline of their distribution in Africa is given. T h e present classification has evolved from earlier classifications, including the so-called 'Yangambi' classification ( C C T A / C S A , 1956; Trochain, 1957; Boughey, 19576, 1961; M o n o d , 1963; Aubréville, 1965; Beard, 1967; GuiUaumet & Koechlin, 1971; Descoings, 1973), and more particularly that of Greenway (1943, 1973; Pratt et al., 1966), but differs in several respects from both. T h e earliest classifications starting with Schimper (1898, 1903) had too few major categories, and failed to m a k e a clear distinction between forest and other w o o d y vegetation types. They also misinterpreted m u c h lowland grassland n o w k n o w n to be edaphic or secondary. T h e Y a n g a m b i classification represents an improvement in some respects, especially in its treatment of forest, woodland and thicket, but there are several reasons w h y it cannot be accepted without considerable modification. T h e main categories are too few to do justice to the variety of African vegetation and they are heavily biased in favour of West African types. T h e most serious criticism, however, concerns the use of the terms 'savanna' and 'steppe' and their definition (for fuller
Classification
discussion see White, in C h a p m a n & White, 1970, and Descoings, 1973, 1978). There is little justification for using the term steppe in tropical Africa. T h e fact that the only ecologist (Walter, 1939, 1943, 1962, 1964) w h o has m a d e a thorough study of the grasslands of eastern Europe and subtropical Africa repudiates the use of the word steppe for the latter should carry great weight. T h e term savanna has been defined in so m a n y different ways that it is n o longer possible to use it in a precise classificatory sense. In more general contexts, however, both popular and scientific, the antithesis 'forest and savanna' is a commonplace and will doubtless continue to be usefully employed for certain tropical landscapes. But its value is greatly diminished if it is m a d e to include, for instance, wooded Sphagnum bogs of temperate regions as is done by Fosberg (1961). Greenway's system is avowedly simple and intended to be understood by laymen. It drew heavily on the experience of m a n y people, and is so well documented that it is possible to attribute actual stands in the field to their correct position. Greenway, w h o avoided the use of imported terms like savanna and steppe, adopted seven main types, namely (1) forest, (2) woodland, (3) wooded grassland, (4) grassland, (5) permanent s w a m p vegetation, (6) bushland, thicket and scrub, and (7) semi-desert vegetation. Because Greenway was only concerned with East Africa it is not surprising that the system adopted here differs from Greenway's in several respects. The chief modifications to Greenway's system are as follows: 1. Mangrove is separated from forest as a major physiognomic type. 2. B a m b o o is treated as a major physiognomic type and not as a type of thicket. 3. Giant-grass thicket is treated as grassland. 4. T h e term scrub is used in a general sense to designate all woody vegetation other than forest, woodland, mangrove and b a m b o o , though in most contexts more precise terms such as bushland or shrubland are preferred. 5. Shrubland is recognized as a major physiognomic type. 6. T h e physiognomically mixed and distinctive Afroalpine vegetation is treated as a major type. 7. Desert is recognized as a major classificatory unit, but semi-desert vegetation is classified as shrubland, grassland, etc., wherever the physiognomy justifies this. 8. T h e physiognomically diverse vegetation of saline and brackish s w a m p is treated collectively as a major classificatory unit. 9. In addition to wooded grassland, three other transitional types, namely scrub forest, transition woodland and scrub woodland are recognized. T h e sixteen major physiognomic divisions of the present classification have been selected and defined so as to correspond as closely as possible to the vegetation on the
45
ground. Thus, forest, woodland, bushland, shrubland, grassland and wooded grassland are not defined arbitrarily, but in such a way as to accommodate the great regional formations of Africa. For the most part the regional vegetation types differ from each other in the height and density of their principal growth forms, but there are often atypical variants in these respects which, however, are typical in most other features. This means that although height and density often have considerable diagnostic value, their application must beflexible,and sometimes they need to be subordinated to other physiognomic features. M a n y applications of this principle are mentioned below where the individual types are described.
Description of the main vegetation types Forest Forest is a continuous stand of trees. The canopy varies in height from 10 to 50 m or more, and usually consists of several layers or storeys. T h e crowns of individual trees interdigitate or overlap each other and are often interlaced with lianes. A shrub layer is normally present. It is usually densest in those types of forest with a more open canopy. The ground layer is often sparse and m a y be absent or consist only of bryophytes. In tropical and subtropical types grasses, if present, are comparatively localized and inconspicuous, though lianes are usually well represented. Epiphytes, including ferns, orchids and large mosses are characteristic of the moister tropical and subtropical types, but vascular epiphytes are virtually absent from more temperate types, if the word temperate is used in a latitudinal rather than a strictly climatic sense. Large epiphytic lichens, especially Usnea, are often conspicuous, especially in upland types. In forest, woody plants, especially trees, contribute most to the physiognomy and the phytomass, and in number of species often greatly exceed herbs. Nearly all the forests in Africa are evergreen or semievergreen, though deciduous forests occur locally. In Madagascar, by contrast, deciduous forest is widely distributed on the drier western side of the island. In m a n y types of forest, especially rain forest, different tree species reach different sizes at maturity, and the overall structure is complex and difficult to analyse. T o overcome this, Davis & Richards (1933-34) adopted the profile diagram, which has been widely used ever since. It is often claimed, and equally often denied, that forests show a well-defined stratification. In recent years a n e w approach to the architecture and growth patterns of rain-forest trees (Hallé & Oldeman, 1970, 1975; Oldeman, 1974; Hallé et al., 1978) and lianes (Cremers, 1973) is greatly enhancing our understanding of structure and stratification, but it has not yet been extensively applied in Africa. In the
46
Regional framework, classification and mapping units
T A B L E 3. Synopsis of the main vegetation types FORMATIONS OF REGIONAL EXTENT:
1. Forest. A continuous stand of trees at least 1 0 m tall, their crowns interlocking. 2. Woodland. A n open stand of trees at least 8 m tall with a canopy cover of 40 per cent or more. The field layer is usually dominated by grasses. 3a. Bushland. A n open stand of bushes usually between 3 and 7 m tall with a canopy cover of 40 per cent or more. 3b. Thicket. A closed stand of bushes and climbers usually between 3 and 7 m tall. 4. Shrubland. A n open or closed stand of shrubs up to 2 m tall.
TRANSITIONAL FORMATIONS OF LOCAL EXTENT:
5. Grassland. Land covered with grasses and other herbs, either without woody plants or the latter not covering more than 10 per cent of the ground. 6. Wooded grassland. Land covered with grasses and other herbs, with woody plants covering between 10 and 40 per cent of the ground. 7. Desert. Arid landscapes with a sparse plant cover, except in depressions where water accumulates. The sandy, stony or rocky substrate contributes more to the appearance of the landscape than does the vegetation. 8. Afroalpine vegetation. Physiognomically mixed vegetation occurring on high mountains where night frosts are liable to occur throughout the year.
present work details of stratification follow the accounts of the original authors. Since some of the terms frequently used to classify forest are applied imprecisely or in conflicting senses the following explanation of the usage adopted here is given: Rain forest. This term is unsatisfactory, both in its connotation and in its application, but is provisionally retained for want of a better. Richards (1952) and others, e.g. Whitmore (1975) and Grubb & Tanner (1976), apply it loosely to include low vegetation, which in the present classification is referred to as scrub forest or thicket. T h e term rain forest seems to be appropriate for the most widespread type of forest in tropical Africa, namely Guineo-Congolian forest on well-drained soils (page 75). T h e upper canopy is nearly everywhere more than 30 m tall. Forest snorter than this occurs very locally in the Guineo-Congohan Region, especially on rocky hills and in upland areas. It is not classified as rain forest, but as short forest, scrub forest or thicket. Guineo-Congolian rain forest includes both evergreen and semi-evergreen variants. Forest which in structure is almost indistinguishable from Guineo-Congohan rain forest and floristically is closely related to it also occurs outside the Guineo-
9. Scrub forest. Intermediate between forest and bushland or thicket. 10. Transition woodland. Intermediate between forest and woodland. 11. Scrub woodland. Stunted woodland less than 8 m tall or vegetation intermediate between woodland and bushland.
EDAPHIC FORMATIONS:
12. Mangrove. Open or closed stands of trees or bushes occurring on shores between highand lowwater mark. Most mangrove species have pneumatophores or are viviparous.
FORMATION OF DISTINCT PHYSIOGNOMY BUT RESTRICTED DISTRIBUTION:
15. Bamboo UNNATURAL VEGETATION:
16. Anthropic landscapes.
13. Herbaceous fresh-water swamp and aquatic vegetation. 13. Halophytic vegetation (saline and brackish swamp).
Congohan Region on the slopes of certain mountains (Chapters VIII and XIII). T h e forests occurring in the lowlands of the wetter eastern half of Madagascar (Chapter X I X ) are somewhat lower in stature than the rain forests on the African mainland but resemble them sufficiently for the term rain forest to be usefully employed. Dry forest. This term is restricted to forests which experience a dry season lasting several months during which atmospheric humidity is low. They are shorter than rain forest, simpler in structure and poorerfloristically.D r y forest is very localized in the Zambezian Region (page 89) and even more so in the Sudanian Region (page 103). It is, however, the most widespread climax type in the West Malagasy Region (Chapter X X ) and occurs on the dry coastal plain of G h a n a (page 176). Semi-evergreen forest. S o m e canopy species are briefly deciduous but not necessarily at the same time. Most members of the understorey are evergreen. The drier types of Guineo-Congohan rain forest are semi-evergreen. It is misleading to call them deciduous. Deciduous forest. The majority of individuals, both in the upper and lower canopy, lose their leaves simultaneously and usually remain bare for several weeks or months. In
Classification
47
universal presence of heliophilous grasses that distinguish woodland from other vegetation types. T h e grasses are usually perennial, but annual grasses are predominant in certain drier transitional types especially under the influence of heavy gracing In most types there is an incomplete understorey of small trees or large Undifferentiated forest. This term is applied to forests bushes, the density of which is very variable. Smaller shrubs also vary greatly in their size and density. Lianes which undergo rapid and kaleidoscopic change in strucare mostly rare or absent. The scarcity of smaller woody ture and composition over short distances. The precise plants is sometimes due to burning of the grassy ground pattern is chiefly of local interest, and in general works, layer, but in some types where the grass cover is sparse, such as this, little subdivision is necessary. Most of the smaller woody plants m a y show an insignificant increase forests in the Afromontane, Zanzibar-Inhambane and after m a n y years of protection from fire. Although the Tongaland-Pondoland Regions are treated as undifgrasses usually dominate the ground layer there is often ferentiated forest. a wealth of herbs and suffrutices, the attractive flowers of which are conspicuous at the end of the dry season With the exception of the sclerophyllous forests of the and early in the rainy season before the grass is tall Mediterranean basin (Chapter VII) most of the imporenough to conceal them. Vascular epiphytes are often tant African forests have been mentioned above. T h e present, though relatively rare, except in secondary forests of the Guinea-Congoha/Zambezia transition woodland occurring on forest sites. zone form a series linking typical Guineo-Congolian forest, through transition woodland, to Zambezian Occasionally, stands of woodland have a closed woodland. A similar transition probably occurred in the canopy and, as a consequence, also a poorly developed Guinea-Congolia/Sudania transition zone, though less grass layer. These are not true forests since they differ evidence survives. from forest in almost all respects other than in density of S w a m p forest andriparianforest are widespread in canopy and field layer. They are best referred to as the Guineo-Congolian, Zambezian, and Sudanian 'closed woodland'. However, when such woodlands conRegions, but in drier regions are rare or are replaced tain forest elements, either because they form part of the there by riparian scrub forest and bushland. In the forest/woodland ecotone or represent a serai stage to Sahara (Chapter XVII) Tamarix very locally forms forest, they are referred to as 'transition woodland' (see riparian forest 10 m high, and Cupressus dupreziana m a y below). formerly have formed forest on the mountains. T h e Woodland as defined above is the most widespread Cape Region is bereft of forest other than enclaves of vegetation type in tropical Africa. It is especially charAfromontane forest and small patches of scrub forest. acteristic of the Sudanian and Zambezian Regions with their continental climate and moderate precipitation, which falls in summer. There is little doubt that m u c h Woodland Sudanian and Zambezian woodland is natural, especially when it occurs on shallow stony soils. However, on Woodland is land with an open stand of trees the crowns deeper soils which have been cultivated, similar woodof which form a canopy from 8 to 20 m or more in land m a y be secondary, having replaced dry forest or height and cover at least 40 per cent of the surface. transition woodland. The crowns of adjacent trees are often in contact but Several of the dominant species of woodland under are not densely interlocking. Frequently the trees are limiting conditions are less than 8 m tall. T h e commore widely spaced and m a y be up to 1 crown diameter munities they form are then scrub woodland. Where apart. In Africa woodlands of regional extent are conscrub woodland occurs in intimate mosaic with normal fined to tropical and subtropical regions. Most African woodland or is part of a gradual continuum it is not woodlands are deciduous or semi-deciduous but nearly given separate treatment. Only a few types which are all types contain a few evergreen species. N o examples more sharply defined are individually described. of completely evergreen woodland have been described from tropical Africa. In woodland the trees (except in In the Somalia-Masai Region, the Sahel transition transitional types) usually have more or less straight zone, and in the Kalahari part of the Kalahari-Highveld boles which are not branched for at least 2 m . If the transition zone only a few trees exceed 8 m in height. boles do branch below this height the branches are The prevalent vegetation is bushland and thicket or usually ascending and progress is unimpeded. T h e various types of wooded grassland. Woodland in these canopy casts little shade and there is usually a ground phytochoria is very local and not very typical; it is not cover consisting principally of herbaceous tussock described separately from the more characteristic types. grasses, the culms of which are up to 2 m high. O n stony Woodland does not occur in the Afromontane or rapidly eroding soils the grass layer is often poorRegion, except when Hagenia abyssinica forms almost ly developed and then other herbs and dwarf shrubs pure semi-open stands with a grassy field layer. are sometimes prominent. It is the dominance of Although forest is the natural vegetation of the trees combined with the light open canopy and almost Mediterranean Region, managed forests, such as some some types the largest trees, as in Zambezian dry deciduous forest (page 90), m a y , on favourable sites or in favourable years, remain evergreen over an almost completely deciduous lower canopy. The West Malagasy forests (Chapter X X ) are deciduous.
48
Regional framework, classification and mapping units
of the Cork O a k (Queráis súber) forests, have the appearance of woodland. Elsewhere, for example in Tetraclinis articúlala communities o n steep rapidly eroding soils, the trees do not always form a closed canopy. The resemblance to woodland, however, is superficial and the community is best regarded as an open forest. Bushland and thicket Bushland is land of which 40 per cent or more is covered by bushes. In this work a bush is defined as a woody plant intermediate in habit between a shrub and a tree. Bushes are usually between 3 and 7 m tall, but can be smaller or larger. They are usually multiple-stemmed and the main axes are frequently 10 c m or more in diameter at the base. Bushy trees are also frequently present in bushland. They too are normally less than 7 m tall and, although there is a main bole, it branches low d o w n so that progress is impeded. Taller trees sometimes occur as émergents but they are either localized in groups or are widely scattered. W h e n they are sufficiently numerous to form a distinct but open canopy w e are dealing with scrub forest. W h e n they are less frequent the term 'wooded bushland' could be used. Grasses are present in most types of bushland but are physiognomically subordinate. Where bushes occur scattered in a continuous sward of grasses and cover less than 40 per cent of the surface w e can talk of 'bushed grassland', but the proportion of bushes is usually m u c h less than this. For reasons given elsewhere (p. 52) bushed grassland is not recognized separately from wooded grassland in this work. Bushland often occurs in rocky or stony places which are unfavourable to grasses, and elsewhere the grasses have been eliminated or m u c h reduced by grazing. Because of the insignificance of grass, it is inappropriate to use the term savanna in such cases although they have frequently been designated as bush- or shrub-savanna in the past. In places where grasses are sparse, the vegetation remains physiognomically bushland even when the cover is m u c h below 40 per cent. Such vegetation can be referred to as open bushland. In thicket, the bushes are so densely interlaced as to form an impenetrable community except along tracks m a d e by animals. In most types of bushland larger or smaller patches of thicket also occur without significant change in floristic composition. Some types of thicket, however, e.g. Itigi thicket (page 97), are dominated by species which do not normally occur in more open communities. Bushland and thicket occur under a wide range of climatic and edaphic conditions which are unfavourable for the growth of taller woody plants. Bushy plants are most widely distributed in Africa in regions where rainfall is between 250 and 500 m m per year, but they only form communities of regional extent where there are two rainy seasons or rainfall is irregular throughout the year, or where dry-season humidity is
very high. Thus, deciduous bushland and thicket is extensively developed in the lowlands of the SomaliaMasai Region (Chapter IV) where in most parts there are two peaks in the annual distribution of rainfall. Elsewhere in Africa, where the annual rainfall is between 250 and 500 m m but falls entirely in summer, grasses are favoured on sandy soils. Consequently, the most widespread vegetation in the Sahel zone (Chapter X V I ) and the Kalahari part of the Kalahari-Highveld transition zone (Chapter X I V ) is wooded grassland, though the woody plants are possibly less dense n o w than formerly because of h u m a n activity. In both areas bushland is largely confined to rocky and stony places, which are relatively infrequent. Deciduous bushland and thicket is also the regional climax vegetation of southwest Madagascar (Chapter X X ) . Here rainfall is between 300 and 500 m m per year and the dry season is u p to 10 months long. Although droughts sometimes last as long as 18 months, rain m a y fall in any month, and relative humidity is high throughout the year. Evergreen and semi-evergreen bushland and thicket occur extensively on the slopes of mountains and other uplands which arise from the lowlands of the SomaliaMasai Region (Chapter IV), and form an ecotone between deciduous bushland and thicket and the drier types of Afromontane forest. M e a n annual rainfall is mostly between 500 and 850 m m and is irregularly distributed throughout the year but with two main peaks. Similar butfloristicallypoorer vegetation also occurs in parts of the Lake Victoria basin (Chapter XII), where the rainfall is somewhat higher (850-1000 m m per year). Evergreen and semi-evergreen bushland is also the characteristic vegetation ofriverbasins in the TongalandPondoland Region (Chapter X V ) . The rainfall there, which is too low to support forest, is concentrated in a single rainy season but there is also significant precipitation in the dry season. In the ZanzibarInhambane Region (Chapter XIII) climatically determined bushland and thicket (other than littoral thicket) is m u c h more localized and occupies only a few pockets of relatively low rainfall. In the Cape Region (Chapter V ) the prevalent vegetation is fynbos. Most fynbos is shrubland, but the tallest examples would be referred to as bushland or thicket if major physiognomic types are defined o n artificial criteria such as height alone. Within the Cape Region, however, there are considerable enclaves in which the climax vegetation was probably true evergreen bushland similar to the drier types of valley bushland of the Tongaland-Pondoland Region (Chapter X V ) . O n most African and Malagasy mountains which are high enough there is a zone of bushland and thicket (Chapters VIII and X I X respectively) above the forest zone. It is usually dominated by Ericaceae. The summits of some mountains which are not high enough to support an Ericaceous belt are covered with elfin thicket. Elfin thicket also crowns a few peaks in the GuineoCongolian Region (Chapter I).
Classification
Evergreen littoral thicket is found on rocky or sandy shores in higher-rainfall areas wherever salt-laden offshore winds are sufficiently severe to prevent the development of forest. It is not included in this work. In tropical Africa the remaining types of bushland and thicket (mostly the latter) are edaphically determined. The most widespread occupy old termite mounds or rocky outcrops. Bushland and thicket, physiognomically similar to that of the Ericaceous belt on the African mountains, occurs locally o n badly drained coastal sands in the Zanzibar-Inhambane Region (Chapter XIII). In the drier parts of the Zambezian Region (Chapter II) dense deciduous thicket occurs on certain soils which favour the intensive root systems of thicket species. There is an abundant supply of water in the rainy season but the soils dry out, at least in their upper layers, in the dry season. In the Maghreb (Chapter VII), forest nearly everywhere represents the climax. It is possible, however, that in certain drier areas, especially on clay soils, bushland and thicket were the original vegetation (page 157). Secondary thicket is also widespread, especially as an early serai stage in the reversion to forest, both lowland and upland. S o m e examples are described by Clayton (1958a, 1961). Just as some communities which are more than 3 m tall, phylogenetically must be regarded as shrubland, other communities less than 3 m in height phylogenetically are bushland, and are treated as such in the following pages.
Shrubland Shrubland is land dominated by shrubs which vary in height from 10 c m to 2 m or more. It occurs where taller w o o d y plants are excluded by low rainfall, summer drought, low temperatures, exposure to wind, or shallowness, salinity, toxicity or extreme oligotrophy of the soil, operating singly or in various combinations. In tropical and subtropical semi-desert areas with summer rainfall the climate is equally favourable to grasses and w o o d y plants. Grasses are dominant on deep sandy soils, and woody plants in stony and rocky places. T h e latter, however, are often bushes or stunted trees, and shrubs play a relatively minor role. The most extensive and distinctive shrublands in Africa are those of the Karoo-Namib Region. Almost the whole of this area, with the exception of the N a m i b desert, is covered with various types of Karoo shrubland consisting almost entirely of shrubs less than 2 m tall. Taller bushes and pachycaul trees also occur locally as scattered émergents but, with the exception of certain types intermediate between Karoo shrubland and Tongaland-Pondoland bushland, they are rarely numerous. T h e Karoo shrublands are markedly different in appearance from the bushlands and thickets of the
49
Somalia-Masai Region, most of which receives only a slightly higher rainfall but lies wholly within the tropics. The dwarf shrublands of the Upper Karoo were formerly separated from the climatic grasslands of the Highveld by a transition zone in which grassy shrubland occurred in mosaic with patches of pure grassland. Overgrazing has largely converted this region into predominantly shrubby communities, and the drier grasslands of the Highveld have suffered a similar fate. T h e original vegetation of the Karoo mountains is believed to have been more grassy than it is today. Overgrazing has favoured the extension of Karoo shrublets, which originally occurred chiefly on shallow soils. The grassiness of these secondary Karoo shrublands varies greatly, and some could more correctly be referred to as shrub grassland, but that distinction is not m a d e here. The very distinctive type of sclerophyllous shrubland k n o w n as fynbos is the prevalent vegetation of the Cape Region. Today most fynbos is less than 2 m tall and only exceptionally more than 3 m , though some types can reach a height of 6 m . T h e scarcity of tall fynbos can partly be explained by the effects of m a n - m a d e fires. Fires, however, also occur naturally, and, in the absence of fire, fynbos becomes moribund and dies, so that tall fynbos was probably never the predominant type in the Cape Region. For this reason and because tall fynbos and the shorter types are part of the same floristic and physiognomic continuum, fynbos is dealt with collectively here—as shrubland. A striking difference between the Cape Region and the Mediterranean Region is that in the former, forest is of restricted occurrence and shrubland (fynbos) is the most widespread climax, whereas in the latter, evergreen sclerophyllous and coniferous forests represent the climax except on the summits of the highest mountains, where dwarf spinous cushion-shaped shrubs prevail. The well-known Mediterranean shrublands, maquis and garrigue, are almost entirely secondary. The tallest types of maquis are somewhat transitional towards bushland, but since they, and less tall variants, often represent part of the same continuous degradation series they are dealt with together as shrubland. Afromontane shrubland is usually grassy and often occurs in mosaic with purer grassland which occurs on shallow soil or along drainage lines. Fire has often favoured the spread of grasses at the expense of the shrubs, so that it is difficult to reconstruct their former relative extent. For this reason they are treated collectively. At higher altitudes, in the Afroalpine belt, dwarf shrubland is a characteristic community, but it is only one component of an extremely diversified assemblage of communities, characterized collectively by the abundance of pachycaul Senecios and Lobelias. Typical shrublands are absent from the GuineoCongolian Region and, with the exception of suffrutex grassland mentioned below, are virtually absent from the Sudanian and Zambezian Regions.
50
Regional framework, classification and mapping units
In the Zambezian Region mixed communities of grasses and geoxylic suffrutices occur extensively on seasonally waterlogged Kalahari Sand and less extensively o n similar soils elsewhere. The suffrutices c o m e into leaf and flower before the onset of the rains while the ground is still bare. A t this time of year the c o m munity is an open dwarf shrubland. The grasses emerge from dormancy two or three months later and eventually, because of their taller growth, completely conceal the subshrubs. Physiognomically the community is n o w a grassland. In some types the phytomass of the subshrubs (especially if their frequently massive underground parts are taken into account) m a y greatly exceed that of the grasses. Since, however, they are physiognomically grassland for a considerable part of the year, and often grade imperceptibly into pure grassland, they are treated, in this work, as grassland. In the lowlands most primary shrubland, with the exception of C a p e fynbos, occurs under a semi-desert climate, where edaphic conditions profoundly influence the vegetation, but are themselves at least in part a consequence of the dry climate. This is true for succulent sub-Mediterranean Euphorbia shrubland, which is largely confined to stony soils, and for Somalia-Masai shrubland occurring on gypseous soils. Halophytic shrubland is described in the chapters o n Desert (XVII) and Azonal Vegetation (XXII). T h e leaves of the shrubland dominants are usually small and are deciduous or evergreen or sometimes reduced to scales. In some types they are sclerophyllous: in others malacophyllous. Shrubs with succulent stems or leaves m a y be present or absent. In some types they are dominant or even almost exclusively present. Grassland Grassland is land covered with grasses and other herbs, with the former physiognomically dominant. In Africa grassland is sometimes completely devoid of w o o d y plants, but such pure grassland is often intimately associated in mosaic or zonally with lightly wooded communities. O n a continental scale, it would be unsatisfactory to attempt to separate them. In this work communities with u p to a 10 per cent cover of w o o d y plants are treated as grassland without further qualification. If the cover of w o o d y plants is between 10 and 40 per cent w e are dealing with wooded grassland which is intermediate between grassland and woodland. Climatically determined wooded grassland is extensively developed in the drier parts of tropical Africa and is dealt with below. Edaphic and secondary wooded grasslands, however, are m u c h more restricted in occurrence, and are often difficult to delimit from the more open grasslands with which they are usually associated. Hence in this work they are treated together. Cyperaceae are present in m a n y types of grassland, particularly edaphic grassland, and locally are more abundant than the grasses themselves, especially in the wettest places. Other herbs, e.g. Acanthaceae, are occa-
sionally dominant. Because of their purely local significance, such non-graminaceous communities are not dealt with separately but are mentioned at the appropriate places in the text. T h e dominant grasses m a y be u p to 3 m or more in height, but are usually shorter. Communities dominated by giant grasses such as Pennisetum purpureum and species of Cymbopogon and Hyparrhenia are regarded as grassland since both in physiognomy and ecology they resemble other grasslands m u c h more than they resemble thicket, to which they are sometimes assigned. Vegetation dominated by bamboos, however, is excluded, and is treated as a separate physiognomic type. S o m e permanent swamps are dominated by grasses. They are described along with other s w a m p communities in Chapter X X I I . Grasses show a remarkable diversity of growth form notwithstanding their basic vegetative uniformity, which is characterized by the almost universal occurrence of a tubular leaf-sheath, an elongate, usually ribbon-like lamina, and long sustained growth of stems and leaves from intercalary meristems. In the wetter parts of tropical and substropical Africa the grasses are generally perennial. Annuals are most frequent in the drier parts and are sometimes dominant. There is, however, n o simple correlation between the distribution of annuals and rainfall. Their extensive dominance is often due to overgrazing. S o m e grasses are procumbent and mat-like and form a dense thin carpet like Cynodon dactylon. M a n y species are tufted, w h e n the culms m a y be virtually leafless {Loudetia simplex) or densely leafy {Hyparrhenia). T h e leaves of some species, e.g. Schismus barbatus, form spreading basal rosettes. Other species, e.g. Imperata cylindrica, have extensive rhizomes which send up solitary or tufted flowering stems. S o m e non-rhizomatous species, such as Schizachyrium platyphyllum, have single culms which are prostrate below and root at the nodes. T h e most conspicuous grasses in the Mediterranean Region and on the high mountains of Africa and Madagascar are tufted species with filiform, sclerophyllous leaves, which can function intermittently throughout the year. S o m e desert and semi-desert grasses are chamaephytes or even nano-phanerophytes. T h e grass flora of Africa is rich and diversified. There are more than a thousand species and they are found in all parts of the continent, but communities dominated by grasses have a very irregular and scattered distribution. S o m e earlier workers such as Schimper (1898, 1903) believed that grassland occurs in tropical Africa as a distinct zonal formation with a distinct type of climate. Schimper distinguished grassland with trees as 'savanna' and grassland without trees as 'steppe'. It is n o w k n o w n , however, that m u c h of the grassland which Schimper and others, e.g. Robyns (1936), thought represented the climatic climax is in fact secondary and is due to fires, mostly m a n - m a d e , or is an edaphic climax due to soil conditions unfavourable to trees. Nevertheless, the
Classification
statement by Richards (1952, p. 316) that 'it is extremely doubtful whether any tropical grassland is a true climatic climax' is possibly too sweeping. According to Walter (1962, 1971) pure grassland without an admixture of woody plants represents the zonal climax vegetation on level areas and average soils in tropical and subtropical regions where the rain falls in s u m m e r and amounts to 100-250 m m per year. Where the rainfall is between 250 and 500 m m per year wooded grassland, which Walter call 'savanna', is the climax. Such grassland and wooded grassland is widespread in the Sahel and Kalahari transition zones. The argument that these 'zonal' wooded grasslands represent edaphic rather than climatic climaxes (see ' W o o d e d Grassland' below) applies equally to pure grassland. The areas occupied by edaphic and secondary grassland vary greatly in different parts of Africa. The most widespread edaphic grasslands are those associated with seasonally or permanently waterlogged soils. They are of limited occurrence in the GuineoCongolian Region, which has a short dry season or none at all. By contrast, they are widespread in the Sudanian, Zambezian and Somalia-Masai Regions and the Indian Ocean Coastal Belt, which all experience strongly seasonal rainfall. Waterlogged soils usually occur in depressions which receive more water than that supplied by the incident rainfall, but the extent of such waterreceiving sites varies greatly from place to place. It depends largely on the stage reached by the landscape in the cycle of geological erosion and on recent geomorphological history. The impeding and reversal of drainage due to warping or tilting of the earth's crust has had a profound effect on the distribution of waterlogged grassland. In some places, parent material has an overriding effect in that waterlogged grassland can occur on eluvial sites such as certain recent volcanic soils (Serengeti Plains, Somalia-Masai Region, Chapter IV), and on soils derived from Karoo mudstone (Luangwa Valley, Zambezian Region). Heavy metal and serpentine soils, which are inimical to the growth of trees in Africa, appear to be almost confined to the Zambezian Region. The great regional vegetation types vary greatly in the ease with which they can be replaced by secondary grassland following h u m a n interference. In the wetter parts of the Guineo-Congolian Region the regeneration of forest after cultivation is so rapid that grasses have difficulty in becoming established. By contrast, the drier peripheral forests are m u c h more vulnerable, and have been replaced by secondary grassland and wooded grassland over extensive areas. Even more vulnerable has been the climax vegetation of the transition zones to the north and south of the Guineo-Congolian Region. Nearly everywhere the original vegetation has gone and the landscape is dominated by fire-maintained grassland. There is evidence that in the Sudanian and Z a m bezian Regions various types of dry forest were formerly more extensive and that they have almost entirely been replaced by secondary grassland or wooded grassland
51
following cultivation and fire. The woodlands in these regions, however, are comprised of fire-tolerant trees and grasses, and have probably always been subjected to at least occasional natural fires. They have withstood the effects of fire and cultivation better than the forests mentioned above. Indeed over large areas agricultural practice is based on the ability of the trees to coppice after felling or lopping of their branches. Experimental evidence shows that m a n y Zambezian woodland trees can survive as annual coppice even when subjected to fierce annual fires for more than forty years. Nevertheless, despite the fire-hardiness of some species, large areas of woodland have been converted to secondary grassland by too intensive farming. In the drier parts of tropical Africa fire is less important. T h e grass growth is not sufficiently luxuriant to support fierce burns and m u c h is removed during the growing season by wild or domesticated animals. Indeed, grazing animals often favour woody plants visà-vis grass by reducing the competitive vigour of the latter. This is also true beyond the tropics. It is believed that the Karoo shrublands were formerly m u c h more grassy than they are today. Within historic times the climax grassland of the drier parts of the Highveld Region in South Africa has been degraded by grazing sheep to secondary Karoo shrubland. Secondary grassland is unimportant in the Cape Region and the wetter parts of the Mediterranean Region, except where deliberately induced for pasture. In some of the drier parts of the Mediterranean Region, however, and in the transition zone between it and the Sahara the landscape is dominated by sclerophyllous grass species, especially Stipa tenacissima. There is increasing evidence that this grassland is secondary and has replaced Pinus halepensis forest. Secondary grassland is n o w the most extensive c o m munity on the African mountains. A s in the lowlands, the vulnerability of the original communities varies greatly. The wettest forests on the wettest mountains are difficult to replace by grassland. At the other extreme the drier forests, especially those dominated by the conifers Juniperus procera and Widdringtonia cupressoides, are very susceptible to fire and can be ignited without being felled. Except for the wettest and driest types, the original vegetation of Madagascar has been devastated by fire and m u c h of the island is n o w covered with secondary grassland. In the present account a clear distinction is m a d e between climatic, edaphic and secondary grassland, but in practice all three factors m a y operate together and it is not always easy to decide to which category a particular type should belong. T h e difficulty over climatic grassland was mentioned above. Elsewhere, grassland m a y occur on soils which are incapable of supporting trees, but the soils themselves m a y have developed under unusual climatic conditions. This m a y be true of the edaphic grasslands of the Accra plains (page 178).
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Regional framework, classification and mapping units
T h e description and classification of African grasslands are in their infancy. For m a n y types even floristic composition has only been sketchily recorded. For the great majority the growth-form spectra and the rhythms of growth in relation to environmental factors and potential competitors remain uninvestigated. Improved classification will depend on a knowledge of the former, and ecological understanding on the latter. The pioneer studies of Walter (1939) on the competitive behaviour of grasses in arid and semi-arid regions, and the publications of Descoings (1971-76) on growth-form analysis, point in the right direction. Descoings, however, recognizes only five main growth-forms, based on the form of the tufts, the manner of branching and the number of culms. This is insufficient since it places together species as different in their ecology (and in other features of growth not used in his classification) as Loudetia simplex and Stipa tenacissima. A more detailed analysis of growth-form would provide a more powerful analytical tool. W o o d e d grassland W o o d e d grassland is land covered with grasses and other herbs, with scattered or, more rarely, grouped w o o d y plants, which are often, but not necessarily, trees. T h e woody plants cover between 10 per cent and 40 per cent of the surface. These figures are somewhat arbitrary, since wooded grassland grades into woodland on the one hand and pure grassland on the other. But by defining wooded grassland in this way, it includes most of the zonal vegetation of the wetter half on the Sahelian Region and of the wetter half of the Kalahari section of the Kalahari-Highveld transition zone. T h e woody plants in wooded grassland, which m a y be trees, bushes, dwarf trees (e.g. Acacia drepanolobium), palm trees or shrubs, are nearly always scattered. W h e n the woody plants are grouped it is better to regard the vegetation as a mosaic. Pure edaphic grassland on seasonally waterlogged soils often forms a mosaic with very sharply defined islands of thicket which occur on the better drained soil of old eroded termite mounds. Such 'termite savanna' is not treated here as a single vegetation type but as a patchwork of edaphic grassland and termite-mound thicket. Greenway (1973) recognizes, in addition to grouped tree grassland, four types of wooded grassland, namely scattered tree grassland, palm-stand grassland, shrub grassland, and dwarf-tree grassland. These categories are certainly useful for local purposes, but extensive areas of wooded grassland often contain trees, bushes and shrubs, and sometimes palm trees, in ever varying proportions, or consist of complicated mosaics of the different variants. For this reason only a single category which embraces all the variants is used in the present work. Climatically determined wooded grassland is described below. Edaphic wooded grassland is also important, especially in the Sudanian and Zambezian Regions,
but since it intergrades with edaphic grassland it is not treated separately from the latter. Secondary wooded grassland is merely an arbitrarily defined phase in a degradation series replacing other vegetation, or in the succession leading tó its restitution. It too is not given separate treatment here. Climatic wooded grassland According to Walter (1971) wooded grassland is the zonal vegetation on deep sandy soils on level sites in tropical and subtropical regions where the rain falls in summer and amounts to 250 to 500 m m per year. H e restricts the term savanna to such vegetation, but, for reasons given elsewhere (Chapman & White, 1970), it is not used in this book except in a more general sense. Where the rainfall exceeds 500 m m per year woodland replaces wooded grassland, and where it is between 250 and 100 m m pure grassland occurs on deep sandy soils. T h e requirements of wooded grassland are found extensively in the southern half of the Sahel zone and in the wetter parts of the Kalahari section of the KalahariHighveld transition zone. It should be pointed out, however, that the sandy soils in both areas are provided by stabilized sand dunes which were formed during the arid phases of the Quaternary. It is doubtful whether similar sandy soils are being actively formed in these transition zones under the present climatic regime. Hence it is somewhat misleading to refer to the wooded grassland as a climatic climax. In the wooded grassland belts, loamy and clayey soils are formed only in periodically wet hollows which are often covered with edaphic grassland. Elsewhere, where there is n o sand cover, a true soil is lacking and the surface is covered with weathered rock. Grasses are not dominant in such places and the vegetation is scrub woodland, bushland or shrubland depending chiefly on the rainfall. It would seem that if the Quaternary sand dunes had not been formed, wooded grassland would be relatively unimportant in the transition zones. T h e vegetation of the Sahel zone has been greatly modified by m a n and his domestic animals, and is certainly less dense than formerly. It is therefore difficult to reconstruct the original vegetation, but comparison with sparsely populated parts of the Kalahari region with a similar rainfall suggests that on sandy soils wooded grassland represents the climax. Desert Deserts and semi-deserts occur in arid regions where plants suffer from lack of water as a result of low rainfall and high evaporation throughout the greater part of the year. The plant cover is sparse and shows various adaptations to unfavourable water conditions. The transition towards arid regions is always gradual unless a range of mountains provides an abrupt climatic division. A n objective criterion for separating arid regions from
Classification
wet regions does not exist and any separation is somewhat arbitrary. Different arid regions have their o w n climatic peculiarities and hence different criteria for their delimitation. In semi-deserts the soil is often more conspicuous than the vegetation, so that the aspect in general view is dominated by the colour and character of the soil rather than by the plants themselves. The latter, however, are still sufficiently evenly distributed and sufficiently numerous for it to be meaningful to refer the c o m munities they form to general physiognomic categories such as semi-desert grassland, semi-desert shrubland, etc. Thus, the grasslands and shrublands of the wetter parts of the K a r o o - N a m i b Region, and the drier parts of the Somalia-Masai Region and the Sahel and Mediterranean-Sahara transition zones comprise semidesert vegetation. In Africa semi-desert usually begins to appear where the m e a n annual rainfall drops to below about 250 m m , but the figure m a y be higher or lower than this, depending o n the distribution of rainfall throughout the year, and its incidence in relation to other factors such as seasonally low temperature, and on soil texture. Desert, like semi-desert, cannot be defined precisely. Very few deserts are so dry that they can be considered 'absolute deserts'. This term applies only to regions where vegetation is completely absent except for oases, or where only ephemeral vegetation develops after a rare rainfall. T h e African deserts comprise the enormous wastes of the Sahara north of the equator, the m u c h smaller but floristically diversified N a m i b desert south of the equator, and some little-known small areas of coastal desert in Somalia. T h e driest parts of northern Kenya, where the rainfall m a y be as low as 150 m m per year, as at Lodwar, west of Lake Turkana, are sometimes regarded as desert. Extensive areas are covered with stone mantles and are almost devoid of vegetation. Because, however, these stone pavements occur in mosaic with more luxuriant vegetation, mostly semi-desert shrubland and dwarf bushland, and the rainfall in most places is higher than at Lodwar, they are considered to be edaphic desert and are not given separate treatment. Following Quézel (1965a), the northern limit of the Sahara desert is shown on the m a p to correspond, more or less, with the 100 m m isohyet, and the southern limit to correspond with the 150 m m isohyet. These criteria, more than any others, are closely correlated with changes in floristic composition which differentiate the flora of the Sahara from those of adjacent areas. Although the vegetation of the moister outer fringes of the Sahara is 'diffuse', in that it is not confined to waterreceiving sites but also occurs on sites with pronounced run-off, the vegetation of the latter is usually sparse, and the contrast between the two types of site is very m u c h more pronounced than in semi-desert areas. With increasing aridity, perennial vegetation becomes more and more contracted until it is eventually confined to
53
water-receiving sites which are so situated that they collect most of the incident rainfall from relatively wide catchment areas. T h e N a m i b desert is not nearly so well differentiated from its surrounding semi-desert vegetation as is the Sahara. Following Walter (1971) the 100 m m isohyet is arbitrarily chosen for its delimitation. The small areas of coastal desert in Somalia are similarly defined. For a number of reasons, most desert vegetation cannot be conveniently accommodated in simple physiognomic classifications. First, some desert plants, e.g. Welwitschia bainesii {mirabilis), have unusual growth forms so that the communities in which they occur d o not fit orthodox categories. Second, vegetation is often too mixed physiognomically, and varies too m u c h and too rapidly from place to place in the relative proportions of its chief physiognomic elements. Such changes are largely dependent o n local variation in the distribution of soil moisture, but are greatly modified by anthropic influences. Third, even when the vegetation is physiognomically uniform, the individual plants, except very locally, and then chiefly in the high mountains, are too few and the vegetation is too exiguous for such terms as 'grassland' or 'shrubland' to be appropriate. Fourth, in some habitats m u c h of the vegetation consists of micro-mosaics of psammophytic, chasmophytic and halophytic elements, the complexity of which renders a simple classification impossible. Since, however, a large proportion of desert species and the communities they form are characteristically associated with distinct physiographic or edaphic features, it is most convenient to classify the vegetation they comprise according to those features. Afroalpine vegetation Afroalpine vegetation, which is confined to the highest mountains of tropical Africa, is physiognomically very mixed. Nearly half the species offloweringplants belong to the following specialized growth forms: giant rosette plants, mostly species of Lobelia and Senecio, up to 6 m tall; tussock grasses and sedges with filiform xeromorphic leaves; acaulescent rosette plants; cushion plants and sclerophyllous shrubs and dwarf shrubs. Owing to the frequency of nocturnal frosts extensive areas of bare soil are subjected to pronounced solifluction giving rise to mobilideserta. In such places unattached colonies of two mosses, Grimmia campestris and G ovalis (ovata), the fruticose lichen, Parmelia sp. near vagans, and one alga, Nostoc commune, are conspicuous. Scrub forest Scrub forest is intermediate in structure between true forest and bushland and thicket. It is normally 10-15 m high. Trees with well-defined and upright boles are usually present but they d o not form a closed canopy; smaller w o o d y plants, principally bushes and shrubs,
54
Regional framework, classification and mapping units
contribute at least as m u c h as the trees to the appearance of the vegetation and its phytomass. In most types of scrub forest in the lowlands of tropical and subtropical Africa, cactiform tree Euphorbias, or more rarely species of Elaeophorbia or arborescent Aloes, are present, but their density varies greatly. In some Euphorbia-dominated scrub forests the tree Euphorbias are said to provide 70-80 per cent cover. Their crowns, however, are very open and the true figure must be m u c h less than this. In types where broadleaved trees are emergent their crown cover is rarely more than 50 per cent and often m u c h less. Also included in scrub forest are a few types which are dominated by bushy trees that rarely have a welldefined bole (and if they do it branches low down). T h e two most important are characterized by Argania spinosa (Mediterranean-Sahara transition zone) and Olea laperrinei (Sahel zone). Both species are often multiple-stemmed but their main axes are of massive proportions. Scrub forest often forms an ecotone between true forest and bushland. O n most of the higher mountains in tropical Africa and Madagascar the stature of the vegetation steadily diminishes with increasing altitude, and scrub forest is sometimes well developed in the transition from forest to montane bushland and thicket. In the lowlands, scrub forest occurs where the rainfall is intermediate between that required by forest and bushland, but it is only well developed where the rainfall is irregular or bimodal in distribution or where dryseason humidity is high. Where there is a single severe dry season and all the rain falls in summer, as in most parts of the Zambezian and Sudanian Regions, scrub forest is very poorly developed, and is not described separately from other vegetation types with which it is associated. Extreme edaphic conditions such as those provided by granite inselbergs and acid peat are also responsible for the occurrence of scrub forest in parts of the Guineo-Congolian Region, though only on a very local scale. Various types of scrub forest are described in the sections dealing with the following phytochoria: GuineoCongolian, Somalia-Masai, Lake Victoria Basin, Zanzibar-Inhambane, Kalahari-Highveld, TongalandPondoland, Sahel and Mediterranean-Sahara transition. Transition woodland The term transition woodland was first used in Nigeria by Jones & Keay (see Forest Department Nigeria, 1948) to designate a type of woodland in which fire-tolerant and fire-sensitive trees occur together. It was originally applied either to secondary, fire-tolerant vegetation occurring on forest sites, which, following a period of fire protection, was reverting to forest, or to degraded forest which had been invaded by fire-tolerant savanna species. Both types are unstable and in the absence of
interference are likely to be transient. This type is described in Chapter I. Keay (1959c) subsequently suggested that the original vegetation of the Guinea-Congolia/Sudania transition zone was a mosaic, with forest in the valley bottom and on the deeper soils of the lower slopes, with Isoberlinia woodland on the shallow soils of the crests, and transition woodland occupying an ecotone between. There is evidence that a similar situation also occurred in the Guinea-Congolia/Zambezia transition zone (Chapter X ) . In the Zanzibar-Inhambane Region (Chapter XIII) Brachystegia spiciformis dominates transition woodland. S o m e stands are clearly serai, but others appear to be stable. There is some evidence that in the Zambezian Region transition woodland represents the climax in an ecotone between dry evergreen forest on the deeper soils and m i o m b o woodland on shallower soils (page 91). Scrub woodland Scrub woodland is intermediate between true woodland and bushland. It is dominated by stunted trees, sometimes no more than 3 m high, which belong to typical woodland species. Scrub woodland sometimes consists almost exclusively of stunted trees but bushy and shrubby species are frequently also abundant. In the Zambezian Region (Chapter II) each of the three main woodland types occurs as scrub woodland under limiting conditions. Thus, scrub m i o m b o dominated by dwarf Brachystegia spiciformis is found towards the upper altitudinal limits of m i o m b o . Other m i o m b o species including B. boehmii dominate scrub woodland on the plateau on unfavourable soils. Throughout the range of mopane (Colophospermum mopane) scrubby communities are found on unfavourable soils, and more locally as a consequence of fire or frost. They also predominate towards the drier climatic limits of the species range. South of the Limpopo River the vegetation of mapping unit 29d, southern undifferentiated Zambezian woodland, is intermediate between Z a m bezian woodland and Tongaland-Pondoland evergreen and semi-evergreen bushland and in places could be regarded as scrub woodland. The ecotone between Zambezian woodland and the edaphic grassland of waterlogged depressions (dambos) is also dominated by scrub woodland, as are some c o m munities on heavy metal and other toxic soils. In this work scrub woodland is not described separately from the other communities with which it is associated. Mangrove Mangrove is dominated by trees or bushes occurring on shores periodically flooded by sea-water. It is sometimes classified as forest but this is misleading since m a n y mangrove species, especially towards their climatic and
Classification
edaphic limits of tolerance, form communities which must be referred to as 'thicket' or 'bushland'. All m a n grove communities, regardless of stature and density, in overall physiognomy resemble each other more than any particular stand of mangrove resembles any other type of vegetation. In a natural classification they should be regarded as a major and quite remarkable physiognomic type. In Africa the tallest mangrove, at the mouth of the Niger River, attains a height of 45 m , whereas at the limits of its geographical range and on unfavourable soils it is no more than 2 m tall. All true mangrove species have either pneumatophores which are exposed at low tide or are viviparous or almost so. Most African species show both these features. Rhizophora has stilt roots which function as pneumatophores. The leaves of mangrove species are thick and leathery. Herbaceous fresh-water swamp and aquatic vegetation Permanent s w a m p , or reed-swamp, occurs in depressions where water accumulates and permanently floods the surface to a shallow depth. In the Guineo-Congolian Region most swampy areas are covered with s w a m p forest, which is described alongside other GuineoCongolian forest. Outside the Guineo-Congolian Region most of the shallower lakes have a wide belt of reed-swamp. Truly aquatic vegetation occurs in deeper water. M a n y s w a m p and aquatic species are distributed through several chorological regions and the vegetation they occur in is classified as azonal (Chapter XXII). The dominants of reed-swamp are usually rooted in the soil and their stemsriseout of the water. The most abundant is the giant sedge, Cyperus papyrus, but other sedges, grasses such as Miscanthus and Phragmites, the bullrush (Typha) and, locally, ferns are also dominant. True aquatics are either completely submerged or have floating leaves. Of the latter, some are rooted in the m u d , whereas others are free-floating. Reed-swamp and aquatic vegetation are often separated by a belt of floating grasses, principally Vossia cuspidata, Paspalidium geminatum and Panicum repens, which is often invaded by the rhizomes of papyrus. T h e vegetation of seasonal swamps is often grassland, which is described in conjunction with other types of grassland. Saline and brackish swamp The vegetation on saline soils, which is dominated by halophytes, is physiognomically varied and includes
55
grassland, wooded grassland, shrubland, and bushland. It is described in Chapter XXII. Most halophytes have fleshy leaves, which in some species are very reduced. The ground between the plants is often covered with a white saline efflorescence. Bamboo Bamboos are giant grasses 2-20 m or more in height, with erect woody stems, which persist for several years. M a n y species flower gregariously over large areas and then die back to the underground rhizomes or die completely. Sometimes they form almost pure, virtually impenetrable communities, but sometimes are scattered in other vegetation. Only four species of b a m b o o (belonging to three genera), namely Arundinaria alpina, A. tesselata, Oreobambos buchwaldii, and Oxytenanthera abyssinica, are indigenous to Africa, though the introduced Bambusa vulgaris is locally naturalized. The b a m b o ofloraof Madagascar is slightly more diversified but less is k n o w n about it. Arundinaria forms Afromontane communities. Oreobambos usually occurs on mountains but is not strictly Afromontane. Oxytenanthera is widespread in the Sudanian and Zambezian Regions. Anthropic landscapes In most parts of Africa the vegetation has been profoundly altered by h u m a n activity and few natural stands remain. T h e only places, however, where the natural vegetation has been totally eliminated over sufficiently extensive areas to show on the m a p are in the Mediterranean Region and the Mediterranean/Sahara Transition Zone, where cultivation has been continuous for more than 2000 years. In other parts of Africa sufficient relics remain to permit plausible reconstruction of the original vegetation. In some of the more densely settled parts of tropical Africa, such as the northern Sudanian Region, cultivation around the big cities is almost continuous. Often the only trees are self-sown individuals of species of economic importance, such as Parkia biglobosa (clappertoniana), Diospyros mespiliformis, Hyphaene thebaica, Acacia albida, Anogeissus leiocarpus and Butyrospermum paradoxum (parkit), which have been permitted to remain, giving the landscape a park-like appearance (wooded farmland).
For reasons given in the Introduction to Part T w o the vegetation of the mapping units is described within a chorological framework rather than under the names of the units themselves. In m a n y instances a single section of text deals only with a single mapping unit, but for other units, especially complex mosaics, the relevant information is given in more than one place. The mapping units are listed in Table 4 (see opposite), with cross-references to the phytochoria under which they are described.
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Part Three
Vegetation of thefloristicregions
Introduction
The vegetation of each of the major phytochoria is described in turn. In addition, three vegetation types, namely mangrove, herbaceous s w a m p and aquatic vegetation, and saline and brackish s w a m p , are included in Chapter XXII, which deals with azonal vegetation.
They are not strictly azonal, since most of their species are confined to the tropics and subtropics. Nevertheless, their ranges transgress the limits of the major phytochoria and it is convenient to give them separate treatment.
i
The African mainland The Guineo-Congolian regional centre of endemism
Geographical position and area
Geographical position and area
Geology and physiography
T h e main area of the Guineo-Congolian Region extends as a broad band north and south of the equator from the Atlantic seaboard eastwards through the Zaire basin to the western slopes of the 'dorsale d u Kivu'. A smaller western satellite occurs in U p p e r Guinea from Guinea Republic to G h a n a . T h e dry ' D a h o m e y interval' separates the two areas. (Area: 2800000 k m 2 . )
Climate Flora Mapping units Vegetation Guineo-Congolian rain forest Hygrophilous coastal evergreen Guineo-Congolian rain forest Mixed moist semi-evergreen Guineo-Congolian rain forest Single-dominant moist evergreen and semi-evergreen Guineo-Congolian rain forest Drier peripheral semi-evergreen Guineo-Congolian rain forest and similar forest in the transition zones Secondary Guineo-Congolian rain forest Pioneer secondary forest Young secondary forest Old secondary forest Guineo-Congolian short forest and scrub forest O n granite inselbergs Upland Parinari excelsa forest in West Africa Guineo-Congolian swamp forest and riparian forest Guineo-Congolian transition woodland Guineo-Congolian elfin thicket Guineo-Congolian edaphic grassland O n hydromorphic soils O n rocky outcrops Guineo-Congolian secondary grassland and wooded grassland Transitional rain forest
Geology and physiography Nearly everywhere the altitude is less than 1000 m but at the eastern end of the Zaire basin the land rises steeply and Guineo-Congolian vegetation gives w a y to Afromontane communities. Between Guinea Republic and G a b o n there are a few limited areas above 1000 m where Guineo-Congolian vegetation is either diluted b y Afromontane species or local upland endemic species, or entirely replaced by Afromontane communities. T h e Zaire basin has an average altitude of 400 m and gradually rises to the uplands and plateaux which form itsrim.In the basin the Precambrian rocks are covered by continental sediments ranging from Palaeozoic to recent. Quaternary sediments cover a 150000 k m 2 alluvial plain in the centre of the basin where there are remnants of former lakes and extensive s w a m p y areas. Surrounding the Zaire basin are the equatorial uplands, a region of dissected plateaux which merge with it. N e a r the basin the plateaux are mostly composed of slightly metamorphic U p p e r Precambrian sandstone, quartzite and schist. T o the east the Zaire basin rises to the Kivu ridge, which consists principally of strongly metamorphic Precambrian gneiss, amphibolite, quartzite and micaschist as well as granitic intrusives. T h e upper slopes lie outside the Guineo-Congolian Region and carry Afromontane vegetation. Towards the north-west the undulating plateau of east C a m e r o u n , which has a n elevation from 600 to 800 m ,risesgently to the B a m e n d a - A d a m a w a highlands which reach 1500 m or more, and consist of basem*nt rocks partly overlain b y volcanic deposits. M o u n t C a m e r o u n , a still active volcano (4095 m ) , stands separate from the main range.
72
Vegetation of thefloristicregions
At the western rim of the Zaire basin the contact with the rather narrow Atlantic coastal plain is formed by the C a m e r o u n - G a b o n plateaux with altitudes of 600-1000 m , which extend south as the Crystal Mts and the M a y o m b e M t s . These ranges consist of gneiss, granite, migmatite, quartzite, greenstone, diorite, micaschist and amphibolite. T h e coastal plain itself, between Angola and Cameroun, varies greatly in width and is especially wide near Libreville where the Ogooue River enters the plain, and also around Douala. The plain is crossed by numerous rivers with mangrove bordering their estuaries. There are also lagoons, lakes and swamps. In West Africa almost the whole of the GuineoCongolian Region is underlain by Precambrian rocks. T h e landscape is formed of relatively low plateaux and plains interrupted by residual inselbergs and small higher plateaux. T h e most important of the latter are Fouta Djalon, the Upper Guinea Highlands ( L o m a M a n 'dorsale') and the Togo-Atacora range, all of which also extend into the Guinea-Congolia/Sudania transition zone. Fouta Djalon is a remarkably level plateau with an average altitude of 1000 m , but ascending locally to 1500 m . It is extensively dissected by the angular drainage pattern. T h e Guinea Highlands attain 1752 m in M t N i m b a and 1947 m in the L o m a M t s . In contrast to Fouta Djalon there are few level surfaces, and the hills are rounded. T h e hardness of the quartzite of the N i m b a chain has helped it to withstand erosion. T h e M a n massif consists of a continuous granite-norite series. T h e Togo-Atacora range rises to over 1000 m in M o u n t Agou. In places along the West African coastal plain between Nigeria and Guinea-Bissau, under the influence of west-east longshore currents, huge sand-bars have formed between the ocean and the lagoons. North of the lagoon stretch the shore is rocky. In Guinea Republic south-west of Fouta Djalon the coastal plain penetrates far inland alongrivervalleys. There are no sand-bars or lagoons but m u d d y creeks and estuaries ('rias') covered with mangrove and flanked by marshes. In GuineaBissau mangrove covers relatively large areas.
Climate Compared with rain-forest areas in other continents, most of the Guineo-Congolian Region is relatively dry and receives between 1600 and 2000 m m of rainfall per year. Areas receiving more rain than this are largely confined to the coastal parts of Upper and Lower Guinea. Only a small part of the Zaire basin receives more than 2000 m m per year. Rainfall in excess of 3000 m m per year falls only in two relatively restricted areas, namely a coastal belt from Guinea Republic to Liberia, but here there is a very pronounced dry season, and a narrow coastal region of Cameroun adjacent to the Gulf
of Biafra. Very locally in the latter at the foot of M t Cameroun annual precipitation exceeds 10000 m m . (See Fig. 5.) The rainfall is, in general, not only lower than in some rain forest regions elsewhere, but its distribution throughout the year is less even. Virtually nowhere in the Guineo-Congolian Region is m e a n monthly rainfall higher than 100 m m throughout the year. In the equatorial parts of the Zaire basin one or two months usually have a rainfall under 100 m m but more than 50 m m . Further away from the equator, but also towards the Atlantic coast at equatorial latitudes, the length and severity of the dry period increases. In the main eastern block of the Guineo-Congolian Region, nearly everywhere the rainfall shows two peaks, separated by one relatively severe and one less severe dry interval. There is a single peak only in the very highrainfall belt of the Gulf of Biafra, which unlike most of tropical Africa lies within the tropical rain belt throughout the year. Individual dry periods are both more frequent and more severe than the climatic diagrams, which are based on m e a n values, indicate. Thus, near the equator in the heart of the Zaire basin periods of several successive rainy days are rather rare and dry periods are frequent throughout the year even during the wettest seasons. At Yangambi, for instance (Bultot, 1954, summarized by Evrard, 1968), dry periods of 6-10 days' duration occur on average 1.6 times per year in the driest month and 0.6 times in the wettest month. D r y periods lasting 30 days or more probably occur once every 12 years. In Nigeria the dry season lasts for three months from December to February, and each receives less than 50 m m . In January and February the harmattan, a north-easterly desiccating wind from the Sahara, at times reaches the rain-forest zone. In Ghana, when the harmattan blows, relative humidity at 15.00 G M T falls to 53 per cent. Further east in Zaire a wind from the same direction also influences the rain-forest climate, but since it emanates from the Ethiopian highlands and the Nile Valley its influence is probably less pronounced than that of the harmattan proper. In G h a n a rainfall varies appreciably over short distances but it is uniformly more than 1750 m m per year in the south-west. Elsewhere this figure is reached only on hills above 600 m elevation. In general there are four to five months with less than 100 m m rainfall, even in the wettest south-west corner. Further west, in Liberia, Sierra Leone and Guinea Republic the rainfall becomes increasingly concentrated in a single season. In Guinea Republic rainfall in places exceeds 4000 m m per year, but for four months there is virtually no rain. According to Aubrévüle (1938) the climax vegetation under this type of climate is not rain forest, because of the severity of the dry season, but there is little published information, and forest of Guineo-Congolian affinity certainly occurs on favourable soils and even extends further to the north-west into a yet more strongly seasonal climate (page 178).
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. 50 m m ) from a wet north-west region with 7-9 wet months. The north-western area has exceptionally high dry-season rainfall, which is an important factor in dry years in the maintenance of the large wildebeest and zebra populations during September and October when the rest of the region is very dry, grass growth has ceased, and m u c h of the grass elsewhere has been burnt. The region dries out from the south-east in M a y to the north-west in July and becomes wetter again from the north-west in August-September to the south-east in December. The annual migrations of wildebeest, zebra, and Thomson's gazelle are closely correlated with this sequence of wetting and drying, as is the timing of grass fires, with the south-western areas burning first in early July. The driest south-eastern areas normally do not burn at all owing to the shortness of the heavily grazed grass. This zone is grazed by the very numerous migratory animals only during the short growing season, and by a few drought-tolerant animals, principally the oryx and Grant's gazelle during the dry season. The commonest large m a m m a l in the Serengeti region is the wildebeest. Its numbers have increased from 263000 in 1961 to 1400000 in 1978. The wildebeest is followed by Thomson's gazelle (c. 400000) and zebra (c. 200000). These three species are predominantly migratory and together constitute c. 60 per cent of the numbers of large m a m m a l s in the region. The remaining large herbivores include buffalo, elephant, Grant's gazelle, topi, hartebeest, and giraffe. In all, twenty-three large herbivores occur in the Serengeti region. Although no integrated account has yet been published of habitat preference and ecological separation, there are indications of the complex relationships which exist. O f the three principal migratory species, the zebra (grazer) selects grass of intermediate length and takes a high proportion of stem and sheath as well as leaf. By contrast, the wildebeest (grazer) is more dependent on green grass and predominantly selects the leaf component of short grass. Thomson's gazelle is a mixed feeder and uses very short grass, frequently after burning or close grazing by wildebeest. A n outstanding feature of the Serengeti ecosystem is the seasonal migrations of the wildebeest, zebra and Thomson's gazelle populations answering the need to seek seasonally available water and grazing. The wildebeest population performs an annual migration, spending the wet season on the Serengeti Plains, then moving westwards towards Lake Victoria at the beginning of the dry season (normally early June), and later northwards to the northern part of the Serengeti Park or into the Masai M a r a G a m e Reserve in Kenya (normally in August or September). W h e n the rain begins again (most frequently in early November) the wildebeest return to the Serengeti Plains, but if the rain is delayed they m a y m o v e south-west first. Variations in the annual pattern
125
of migration can be correlated with differences in rainfall distribution. Like the wildebeest, the zebra and Thomson's gazelle occupy the Serengeti Plains in the rains, and the woody communities to the west and north during the dry season. The zebra usually follows the whole range of movement of the wildebeest, but tends to occupy areas of taller grassland, and the two species only occasionally occupy the same areas at the same time. The Thomson's gazelle population also migrates over the same range as the wildebeest, but does not m o v e as far north. It has been assumed that the animal populations are subject to natural regulation so that they remain within the long-term carrying capacity of their habitats. T h e precise means by which this is attained, however, is far from clear. Despite the incredible increase in size of the wildebeest population during the last twenty years there is n o indication that it is approaching the equilibrium level determined by prevailing climatic conditions. However, a reduction in grass productivity through drought would be expected to result in increased mortality and, depending on the severity of the drought and the population size at the time, this might amount to a population 'crash'. Since this section was written an important synthesis has been published (Sinclair & Norton-Griffiths, eds., 1979). Main vegetation types The grasslands of the Serengeti Plains have been described by Anderson & Talbot (1965) in relation to the distinctive plains soils. Herlocker (1975) has written a synopsis of the woody vegetation of the remainder of the Serengeti National Park, and Glover and T r u m p (1970) have prepared a detailed account of the vegetation of the Narok District of Kenya Masailand which includes the northern fringe of the Serengeti ecosystem. The vegetation of the Ngorongoro Conservation Area is dealt with by Herlocker & Dirschl (1972) in a richly illustrated account. The main vegetation types recorded by these authors are summarized below. The Serengeti ecosystem lies almost entirely within the Somalia-Masai Region but has some distinctive features. T h e grasslands of the Serengeti Plain are unique, and the wooded grasslands to the north and west are different from most other Somalia-Masai y4cûc/'ii-dominated communities, principally in the insignificance of bushy plants other than Acacia and Commiphora and the relative abundance of grasses, especially perennial species. The extent to which these features might be due to the prevalent grass fires and a large ungulate population is uncertain. 1. Edaphic grassland of the Serengeti Plains The grasslands of the Serengeti Plains grow on soils derived from volcanic ash and are briefly mentioned on page 116 in relation to other edaphic grasslands of the
126
Vegetation of the floristic regions
S o m a l i a - M a s a i Region. Grasslands occurring o n volcanic ash are of very restricted occurrence in Africa a n d are almost confined to the Serengeti region, where the ash w a s supplied b y t w o volcanoes, o n e extinct, the other active. This unique pedological feature m a y provide the explanation for the unique concentration of large m a m m a l s in this part of Africa (page 127). T h e extinct volcano, Kerimasi, at the northern end of the Crater Highlands, in the last stages of eruption, c. 150000 B.P., produced huge quantities of fine whitishgrey ash which included a high proportion of calcium carbonate. T h e ash fell over a wide area, including the south-eastern Serengeti, where it settled in successive layers, filling the hollows a n d providing a relatively flat surface to the formerly undulating peneplain. It hardened to form grey a n d light-brown calcareous tuffs, and, with the accumulation of lime through d o w n w a r d leaching, almost continuous layers of calcitic hard-pan formed at several successive horizons. T h e present surface of the eastern plains of the Serengeti consists of grey, dusty, easily erodible soil derived from the upper layers of the tuff a n d is precariously protected from the erosive effects of wind, rain, a n d animals b y the short plant cover. A s the activity of Kerimasi ended, a n e w volcano, Oldoinyo Lengai, arose 11 k m to the north, a n d is n o w the only active volcano in the region. T h e last eruption w a s in 1967. Its blackish-brown ash has locally formed tuffs a n d agglomerates in the eastern Serengeti but m u c h has remained unconsolidated as loose black sand, which forms elongated dunes m o v i n g in a south-westerly direction. T h e Serengeti Plains, which lie partly inside partly outside the Serengeti National Park, cover s o m e 6 2 5 0 k m 2 of gently rolling grassland between 1350 a n d 1 6 5 0 m above sea-level. M o r e than fifty species of grass are c o m m o n a n d a further fifty species also occur. A t certain times of year the grasslands provide pasture for the world's greatest concentration of large m a m m a l s , estimated at m o r e than 1 million individuals a n d belonging to thirty species of ungulate, a n d eight species of large predator. Rainfall gradually increases from 380 m m per year in the east to 780 m m in the west. T h u s , the gradient from the juvenile ash soils in the east to the m o r e mature b r o w n calcareous soils of the west is paralleled b y a climatic gradient. A n d e r s o n & Talbot (1965), w h o have described zonation of soils a n d vegetation, a n d indirectly the distribution of the large ungulates, in relation to these gradients, recognize six types of grassland, although the change is gradual. It seems that the soils derived from volcanic ash, especially vertisols, at least under the present climatic regime, favour grass rather than w o o d y vegetation, whereas o n the granite soils in the higher-rainfall areas to the west, trees are excluded, at least in part, b y fire, though the matter is b y n o m e a n s fully elucidated. T h e m a i n features of A n d e r s o n & Talbot's grassland types are as follows:
1. Sparse grassland on juvenile soils on volcanic ash. T h e ash forms both mobile a n d stable dunes. Rainfall is 3 8 0 - 5 0 0 m m per year, but, because of high porosity a n d low moisture retentivity of the soils, the area is edaphic desert. T h e soils are fully saturated with bases but salt accumulation is very local; p H is 7.4 near the surface a n d 7.9 at 110 c m ; at 200 c m , in the subsoil of the previous soil surface, it reaches 9.7. T h e vegetation is sparse with about 1 5 - 2 0 per cent basal cover. A m o n g the early colonizers Chloris gayana, Dactyloctenium sp., Digitaria macroblephara, Sporobolus ioclados a n d S. kentrophyllus are c o m m o n . O n m o r e stable areas between the dunes scattered bushes of Acacia mellifera sometimes occur. 2. Short grassland on calcimorphic soils with hardpans. This type covers a large part of the Eastern Serengeti Plains. T h e parent material is fine volcanic dust overlying calcareous tuff. A very hard calcareous p a n occurs at a depth of about 95 c m . T h e soils are base-saturated a n d have a high exchangeable sodium content; p H increases from 8.1 at 15 c m to 9.8 at 100 c m . T h e vegetation is short a n d sparse (basal cover ± 20 per cent). T h e m o s t characteristic species are a sedge (Kyllinga) a n d a dozen species of Sporobolus. Inside the Park this area is heavily grazed b y wild ungulates during the wet season, a n d outside b y domestic animals. T h e soils are very friable a n d vulnerable to w i n d erosion w h e n the vegetative cover has been d a m a g e d , as it has been in parts of the N g o r o n g o r o Conservation A r e a b y excessive livestock. 3. Intermediate grassland on calcimorphic soils with soft pans. This type is intermediate between grassland type 2 a n d taller grasslands to the west. T h e soils are not quite fully base-saturated above, but are completely so below. T h e upper layers are salt-free a n d p H increases from 6.2 near the surface to 9.1 at 100 c m . T h e taller species are Pennisetum mezianum, Eragrostis tenuifolia and Sporobolus spp. with lower patches of Andropogon greenwayi, Panicum coloratum, Cynodon dactylon, etc. Basal cover is c. 30 per cent. This area is heavily grazed but occasionally accumulates sufficient fuel to burn. 4. Tall grassland on vertisols of lithom*orphic origin. These soils are derived from fine ash overlying tuff; p H is 6.8 near the surface a n d 7.4 at 100 c m . T h e y have a m u c h deeper rooting zone than those described above. There is a high degree of base saturation but an absence of soluble salts to a depth of 170 c m . There are concretions of calcium carbonate but there is n o pan. T h e most abundant species are Andropogon greenwayi, Digitaria macroblephara, Cynodon dactylon, Eustachys paspaloides, with Themeda triandra, Pennisetum stramineum, P . mezianum a n d Michrochloa kunthii also present. Basal cover averages 50 per cent. These grasslands are only grazed sporadically b y the herds of animals. T h e latter avoid heavy-textured soils w h e n they are wet, and m o v e eastwards where m a n y of the s a m e plants are available
The Somalia-Masai regional centre ofendemism
in shorter and apparently more palatable forms. Most of this grassland is burned at least once annually.
111
Andropogon greenwayi, begins late, this vegetation is acceptable for a longer period. The tall grassland offers good forage at the beginning of the wet season, but as it 5. Intermediate grassland on southern vertisols of litho- is not normally grazed frequently during the growing morphic origin. T h e soils are somewhat similar to the season, its quality rapidly declines. northern vertisols but appear to be derived solely from During dry periods in the wet season the short grasscalcareous tuff, and are not fully saturated with bases lands are the first to become unproductive. T h e interuntil a depth of 100 c m . T h e dominant grasses are mediate and tall grasslands continue to produce for a Pennisetum mezianum, P. stramineum, Cynodon dactylon longer time and the grazing animals use them as a and Andropogon greenwayi. temporary food supply until the short grasslands receive rain again. During the dry season, the Serengeti Plains 6. Tall grassland on brown calcareous soils. These lighter- as a whole receive very Utile rainfall, forage production virtually ceases, and water and shade are virtually textured soils occur in the extreme west and along the absent. T h e animals consequently m o v e to the woodnorthern edge of the plains. The parent material is a lands where conditions are more favourable. calcareous conglomerate with some quartzitic gravel, originating from the decomposition of granite at and Various reasons have been offered to explain w h y the near the surface. There is negligible ash deposition. The ungulates prefer the shorter grasslands. Anderson soils are m u c h better drained than the vertisols and have & Talbot (1965) believed that the animals avoid the lower base status and p H (6.2 near the surface but 8.6 at sticky soils of the taller grasslands and prefer certain 130 c m ) . Soluble salts are present in the less-permeable growth forms and developmental stages of the shorter lower layers. T h e dominant grass is Themeda triandra, grasses. Bell suggested that grass height is an important closely followed by Pennisetum mezianum and P. stramifactor in relation to feeding adaptations and also that nium. Basal cover averages 45 per cent. T h e grasslands the hazard of prédation is lower in short grassland. are heavily grazed in relatively dry years, but are only Kreulen (pers. c o m m . in Lamprey, 1979) considers lightly grazed in wet years. They are normally burned that the main reason is the availability in the short grassannually. lands of essential nutrients to meet the high requirements of lactation during the months February-May, the first four months of growth of wildebeest calves. H e In the wooded areas of the Serengeti ecosystem very observed that the areas of highest occupance during gentle slopes towards the bottom of the catena have lactation are those in which the grasses and the water alkaline soils with impeded drainage which support have a relatively high calcium content, which is necesgrassland, but their area is very m u c h less than that of sary for milk production without loss of condition in the the w o o d y communities. mother. H e also found that Themeda triandra, the Subsequent to the publication of Anderson & predominant grass in the tall western grasslands, conTalbot's paper m u c h research, mostly unpublished, has tains too little calcium to enable the lactating females to been done on the Serengeti grasslands and is summaintain a positive calcium balance. marized by Lamprey (1979). Fire-control experiments Hence it appears that: 'The unique concentration of indicate that the Themeda grassland of the western ungulate animals (primarily wildebeest) which use the Serengeti Plains is maintained mainly by the fires, which eastern Serengeti Plains m a y depend upon the equally occur at least once every three years. They also show unique occurrence of the highly calcific ash soils that Acacia tortHis will only regenerate if fire does not produced by the eruption of the n o w extinct volcano, occur for five consecutive years. In the absence of fire Kerimasi' (Lamprey, 1979). most Acacia species grow at c. 1 m/year and become tolerant to moderate fires after 3-4 years. Where giraffe are numerous their browsing offtake m a y nearly equal 2. Secondary grassland of the Loita Plains the growth for m a n y years and render the regenerating trees vulnerable to burning for an extended time. The Loita and similar contiguous plains occur in the Narok district of Kenya Masailand 90 k m north of the W h e n the short grass plains are fenced to exclude northern fringe of the Serengeti Plains. In some ways the animals they grow a dense grass cover 40-50 c m high, Loita grasslands are similar to those of the Serengeti while the surrounding grassland remains at less than Plains and are part of the same ecosystem, but their 10 c m high owing to ungulate grazing. This taller sward origin is different, since they occur on truncated soils is easy to burn. After burning there is an increase in and have replaced evergreen bushland, following its density of the fire-tolerant species Themeda triandra, degradation by fire and browsing. which is normally scarce or absent on the eastern plains. The Loita plains he roughly between 1 ° S. and 2° S. The different types of grassland vary greatly in the and 35° E . and 36° E . , between 1700 and 1900 m , and quality of their grazing. The short grassland provides the cover an area of 4500 k m 2 . In the north and west they lie best herbage, but as the rainfall is low and irregular, this on Tertiary-Recent volcanic rocks and in the south on good forage is available for only a short time. The interthe Basem*nt Complex. The volcanic rocks are phonomediate grassland has a longer growing period and lites with thin intercalated tuffs. T h e soils to the north since the growth of stem in the predominant species,
128
Vegetation of the floristic regions
and east are composed of lava dust and sediments forming brown calcareous loams and grey, compacted, loamy sands. T o the south they are shallow and stony with rock outcrops and large areas of black clay. All these soils are severely truncated. T h e rainy season lasts from N o v e m b e r to June and m e a n annual rainfall ranges from 1000 m m in the north and west to 500 m m in the extreme east towards the edge of the Rift Valley. T h e plains are grazed by Thomson's gazelle, Grant's gazelle and kongoni, and by large migratory herds of wildebeest, zebra, and topi. Except for small fringe areas to the south and west, inhabited by tsetse flies, the whole of the plains are heavily grazed throughout the year by Masai cattle, sheep and goats. Glover & T r u m p (1970) recognize two kinds of grassland, tall and short. Tall grassland is 45 c m to 2 m tall and is composed principally of Pennisetum mezianum, P. schimperi, Hyparrhenia cymbaria, H . filipéndula, H . hirta, Hyperthelia dissoluta, Themeda triandra and Dichanthium insculptum. Short grassland is closely cropped vegetation often no more than 7 c m tall, but there m a y be some flowering culms up to 45 c m or more. It usually occurs on shallow or compacted soils in severely overgrazed and trampled areas. T h e principal species are Microchloa kunthii, Sporobolus festivus, Cynodonon dactylon and some constituents of the tall grass zones such as Themeda and Dichanthium insculptum. T h e short grasslands are favoured by m a n y wild ungulates and Masai sheep and goats because they produce fresh green shoots throughout the year. The distribution of tall and short grassland is often controlled by soil depth and micro-relief associated with termite m o u n d s (Glover et al., 1964) and erosion terraces (Glover & Wateridge, 1968).
dichogamus, Euphorbia candelabrum, E. nyikae, E. tirucalli, Salvadora pérsica and Sansevieria ehrenbergii. 4. Acacia-Commiphora deciduous wooded grassland and related types
This type is classified by Herlocker as woodland. In most places, however, the canopy cover is less than 40 per cent and the trees are only 4-7 m high, so that the vegetation fits the category of wooded grassland as defined in this work (page 52). Acacia-Commiphora wooded grassland is the most extensive w o o d y vegetation type in the Serengeti National Park and covers 7260 k m 2 or 88 per cent of all w o o d y vegetation. It consists of a single open stratum of Acacia or Commiphora thorn-trees mostly from 3 to 7 m high but in a few species from 9 to 20 m . Shrubs and bushes are poorly represented, but scattered single bushes or small groups of Grewia fallax and Cordia ovalis sometimes form a very open understorey. There is a characteristic grass stratum 0.5-1.5 m high. It is dominated by such species as Digitada macroblephara, Themeda triandra and Eustachys paspaloides on relatively well-drained soils, and by Pennisetum mezianum on poorly drained soils. Herlocker, for mapping purposes, divides this vegetation type into 39 species types, 38 of which have one or more of 11 species of Acacia (A. nilotica, A. hockii, A. Senegal, A. gerrardii, A. robusta subsp. usambarensis, A. drepanolobium, A. seyal, A. xanthophloea, A. sieberana, A. tortilis, A. polyacantha) as dominants or co-dominants. Commiphora schimperi is the sole dominant of the remaining species type. Most of these species combine and recombine in a kaleidoscopic fashion and, as little is k n o w n of their ecology, interpretation of the species types is sometimes difficult. There are, however, some distinctive variants. Acacia gerrardii occupies large areas of poorly 3. Acacia-Commiphora deciduous bushland and thicket drained clay soils. It also dominates secondary wooded grassland replacing evergreen bushland. This type, the most extensive and characteristic of the Acacia drepanolobium often forms almost pure 1-8 m Somalia-Masai Region, is very poorly represented inside high open stands o n poorly drained soils of valley the Serengeti National Park. In most places the rainfall bottoms. It is c o m m o n to find stands of dead trees, is too high; elsewhere it is precluded by the volcanic ash suggesting recent extreme variation in edaphic consoils of the Serengeti Plains. It is, however, well develditions. oped outside the Park along the drier eastern fringe of the Serengeti ecosystem. Commiphora schimperi is dominant on well-drained soils derived from granite or granitic gneiss onridgetops Inside the Park it is dominated by 2-6 m high Acacia and slopes. Its most important associates are Acacia mellifera and occurs o n termite m o u n d s , and in distortilis, A. robusta subsp. usambarensis, A. Senegal and A. turbed places where bare soil is associated with sheet hockii. and gully erosion or salt-licks. Acacia tortilis is most frequently dominant in the Outside the Park it is widespread in the Ngorongoro drier eastern part of the Park, where it forms 9-14 m Conservation Area, e.g. in the Doinyoogol Hills high stands on the fringes of the Serengeti Plains. There (Herlocker & Dirschl, phot. p. 16), o n the Lake Eyasi is little regeneration and the stands are degenerating escarpment andflats(Herlocker & Dirschl, phots, p. 27) because of the death of old trees. and in Oldupai Gorge (Herlocker & Dirschl, phots, p. 18). Characteristic species include: Acacia drepanoloAcacia robusta subsp. usambarensis is most bium, A. mellifera, A. seyal, A. tortilis, Adansonia digitata,frequently dominant in the wetter, western half of the Cissus cactiformis, C. quadrannularis, Commiphora park where it often forms almost pure, even-aged stands madagascariensis, C. merkeri, Cordia sinensis, Crotón 8-12 m high with dense canopies and little regeneration.
The Somalia-Masai regional centre ofendemism
Acacia xanthophloea, which reaches a height of 22 m , is a riparian species, or otherwise dependent on ground water. Acacia sieberana and A. polyacantha are also riparian species in this part of their geographical range. They are not typical Somalia-Masai species but are more characteristic of regions with higher rainfall.
129
harveyi, Cassine buchananii, Euclea divinorum, Lannea stuhlmannii, Olea africana, Pappea capensis and Ziziphus mucronata. Succulents include Euphorbia candelabrum, Aloe volkensii and species of Sansevieria and Kalanchoe. 7. Evergreen forest
T h e total area occupied by the surviving remnants of evergreen forest and evergreen bushland with which it is 5. Combretum-Terminalia secondary wooded grassland usually associated is small, amounting to no more than This type is a fire-climax which has replaced dry ever240 k m 2 . green forest o n ridge tops and upper slopes in the Evergreen forest occurs on alluvial soils as narrow northern part of the Park. It occupies c. 500 k m 2 . T h e and often discontinuous riverine communities in the open overstorey is dominated by Combretum molle and M a r a River basin and along the lower Grumeti, Orangi Terminalia mollis, which grow to 10-13 and 15-17 m in and Mbalageti rivers. Towards the north of the Park height, respectively. T h e open understorey is dominated there are also small relict patches of dry evergreen forest by Heeria reticulata, Acacia nilotica subsp. subalata, and and scrub forest on the deep sandy loams of broad ridge A. hockii. T h e 1-2 m high grass stratum, which burns tops. In this situation they are usually associated with fiercely every dry season, is dominated by species of evergreen bushland on shallower soils. These two types Diheteropogon, Hyparrhenia, Loudetia and Themeda. T h e were probably dominant north of the Grumeti-Mara principal species on termite m o u n d s are Rhus natalensis River divide. Their area has been greatly reduced by fire, and Grewia trichocarpa with emergent Lannea stuhland they have been extensively replaced by Combretum mannii and Sclerocarya birrea which reach the main molle, Terminalia mollis wooded grassland and Acacia canopy. Parinari curatellifolia is characteristic of termite robusta subsp. clavigera, A. gerrardii wooded grassland m o u n d s along seepage lines. It is believed that a signirespectively. ficant decrease in the Combretum and Terminalia T h e canopy of riparian forest is composed princippopulations has taken place in recent years because of ally of Aphonia senegalensis, Ekebergia capensis, Ficus elephants, but that large trees of Terminalia have sufspp., Garcinia livingstonei, Lecaniodiscus fraxinifolius, fered least, because they can withstand pushing over. Tamarindus indica and Ziziphus pubescens. Both Combretum molle and Acacia hockii regenerate In rain-fed dry evergreen forest the main canopy freely from their roots and persist despite repeated constituents are Diospyros abyssinica, Drypetes gerrardii, burning, which, however, retards their further developCassine buchananii, Lecaniodiscus fraxinifolius, Suregada ment. procera and Teclea nobilis, with Chaetacme aristata, Euclea divinorum, Olea africana, and Schrebera alata occurring less frequently. Capparis erythrocarpos, Crotón 6. Evergreen and semi-evergreen bushland and thicket dichogamus and Teclea trichocarpa are the most abundant m e m b e r s of the understorey. T h e broadThis type has a scattered distribution, chiefly on stream leaved grass Setaria chevalieri occurs in the field layer. banks, rocky hills, along seepage lines and on termite m o u n d s , throughout most of the Park, but is best developed in the north, where, however, is has been exten8. Afromontane communities sively destroyed by fire (see 7 below). O n rocky hills near the K e n y a border it is dominated by Euclea raceT h e Crater Highlands are outside the Serengeti mosa subsp. schimperi, Haplocoelum foliolosum, Tarenna ecosystem but within the greater Serengeti region. They graveolens, Teclea nobilis, and T. trichocarpa, associated rise from Lake Eyasi at 1000 m to an extensive high with Aloe, sp., Cordia ovalis, Euphorbia candelabrum, plateau with an average elevation of 2150-2450 m . Grewia trichocarpa, Pappea capensis, Rhus natalensis and Several extinct volcanoes, of which the highest is Strychnos henningsii. Lolmalasin Mountain (3350 m ) , rise above the plateau and several calderas, including Ngorongoro, are sunk Evergreen bushland is the climax throughout m u c h into it. T h e Crater Highlands are drier than most massifs of south Narok District in K e n y a Masailand i m m e in Africa of comparable size and altitude, and their diately north of the Park, but very little primary or Afromontane communities show several distinctive undisturbed vegetation remains, and nearly everywhere features. Also, because of the complex physiography and it has been replaced by secondary communities, includheavy grazing of the vegetation by wild ungulates and ing the severely degraded secondary grasslands of the domestic stock, Afromontane vegetation is sometimes Loita Plains (see above). T h e principal dominants are Acacia brevispica (on shallow, stony soils), Carissa edulis, less sharply differentiated from 'lowland' vegetation than is often the case. Herlocker & Dirschl (1972) Crotón dichogamus, Grewia similis, Osyris sp., Rhus recognize the following main types: natalensis, Tarenna graveolens and Teclea simplicifo— Artemisia afra, Erica arbórea montane heath above lia. O n deeper soils and along dry watercourses the 2450 m. following occur as stunted emergent trees: Albizia
130
Vegetation of thefloristicregions
— Crotón macrostachyus, Calodendrum capense, Olea spp., Albizia gummifera montane forest above 2450 m . — Vernonia auriculifera, Crotalaria agatiflora subsp. imperialis secondary thicket above 2450 m . — Hagenia abyssinica, Gnidia glauca upper m o n t a n e forest, above 2700 m .
— Juniperus procera dry evergreen forest in steep canyons between 2 4 5 0 and 2900 m . — Eleusine jaegeri, Pennisetum schimperi m o n t a n e grassland, c. 2300 m . — Acacia lahai woodland, which is probably secondary, 2100-2450 m . — Arundinaria alpina b a m b o o , above 2300 m .
v
The Cape regional centre of endemism
Geographical position and area Geology and physiography Climate Flora Mapping unit Vegetation Cape shrubland (fynbos) Secondary Cape shrubland (Rhenosterbosveld) Coastal bushland and thicket Riparian bushland and thicket The transition to Karoo
Geographical position and area This region covers the south-western and southern part of the Cape Province, South Africa, between 32° and 35° S. and 18° and 27° E. Typical Cape vegetation does not occupy the whole of this area. There are large enclaves of Karoo and Afromontane vegetation and small patches of bushland of Tongaland-Pondoland affinity. The easternmost extremity of Cape vegetation in the Suurberge is separated from the main block by the bushlands of the Sundays River valley. Outliers of Cape vegetation to the north occupy the highlands above Van Rhynsdorp and the summit of the Kamiesberg. (Area: 71000 km 2 .)
Geology and physiography The landscape is dominated by subparallel folded mountain ranges with an average altitude of 10001500 m and individual peaks ^exceeding 2000 m . T h e major ranges are constructed of Table Mountain sandstone and the minor ones of smaller sandstone folds or of Witteberg quartzite. In the western part the foothills and lower slopes are commonly formed of Cape granite. The valleys and parts of the coast belt are formed from Bokkeveld shales and sandstones of the Cape System and Malmesbury shales of the late Precambrian. T h e coastal fringe itself consists of Tertiary to Recent sands, conglomerate, and limestone.
Climate Rainfall exceeds 250 m m per year, and is mostly from 300 to 2500 m m , but locally in the mountains reaches 5000 m m . The western part receives 60-80 per cent of its rain in winter but from Swellendam eastwards the rainfall is more evenly distributed throughout the year. O n the higher mountains summer drought is alleviated by moisture-bearing clouds from the south-east, and on the west coast sea mists are fairly frequent. During winter, snow falls regularly on the higher mountains, especially in the west, but persists only on the southern slopes. In general the winters are mild. Frosts are unknown at the coast, but occur inland and in the mountains where they
132
Vegetation of thefloristicregions
are infrequent or not severe. Strong desiccating winds blow at certain seasons. (See Fig. 11.)
Flora There are about 7000 species, of which more than half are endemic. Endemic families. Bruniaceae (12 genera, 75 species). Geissolomataceae (1 species). Grubbiaceae (2 genera, 5 species). Penaeaceae (5 genera, 25 species). Retziaceae (1 species). Roridulaceae (1 genus, 2 species). Stilbaceae (5 genera, 12 species). Other characteristic families. Ericaceae (18 endemic genera and c. 650 endemic species). Proteaceae (11 endemic genera and c. 320 endemic species). Restionaceae (c. 10 endemic genera and c. 180 endemic species). Rutaceae-Diosmeae (10 endemic genera and c. 150 endemic species). Endemic genera. About 210 genera are confined to the Cape Region and a further 70 have their greatest concentration of species there. A m o n g the latter are Agathosma (130 endemic species), Aspalathus (240), Cliffortia (70), Crassula (145), Erica (520), Ficinia (50), Metalasia (30), Muraltia (100), Phylica (140), Protea (85), and Restio (40). Most of these genera are virtually confined to the Cape Region. Erica, however, has about 35 species on the mountains of tropical Africa and in the Holarctic Kingdom, and Protea has about 40 species in tropical Africa.
Mapping unit 50. Cape shrubland (fynbos). Only a single unit appears on the m a p , but as is shown below, the typical Cape vegetation is modified in places by the occurrence of species of karroid and tropical affinity.
Vegetation T h e prevalent vegetation of the Cape Region is fynbos, which most characteristically occurs in the form of 1-3 m tall sclerophyllous shrubland. Large parts of the Cape lowlands, however, where they are not cultivated, are today occupied by secondary shrubland dominated by the 'rhenosterbos', Elytropappus rhinocerotis. There is some evidence that the original vegetation, which has been replaced by Elytropappus, included m a n y species of tropical and karroid affinity. O n the coastal plain itself coastal fynbos is the prevalent vegetation, but locally there are patches of bushland and thicket dominated mainly by tropical species. Mountain streams in m a n y parts of the Cape Region are fringed with riparian
thicket and scrub forest dominated by a mixture of Cape endemics and Afromontane species. T h e term fynbos is applied to virtually all the terrestrial vegetation of the Cape Region other than the enclaves mentioned above. Despite its wide range of variation in floristics and structure, most fynbos fits the definition of shrubland or bushy shrubland adopted in this work. Only a few communities of specialized or otherwise localized habitats belong to other physiognomic types. T h e Protea bushes and other tall plants that often occur scattered in bushy fynbos m a y thicken up and form dense impenetrable thickets 4-6 m high if they receive sufficiently long protection from fire. Experiments have shown, however (F. J. Kruger, pers. c o m m . ) , that most fynbos species cannot regenerate under these conditions and become moribund and die. Accordingly, such thickets, even in the absence of fire, represent an impermanent phenomenon, and are not described separately below. Cape shrubland (fynbos) (mapping unit 50) Refs.: Aco*cks (1975, p. 104-7); Adamson (1927; 1938a, p. 8695); Duthie (1929); Marloth (1908); H . C . Taylor (19636; 1972a; 19726; 1978); Wergere/a/. (1972). Phots.: Aco*cks (1975: 102, 103, 104); Adamson (1927: 3-8; 1938a; 1, 2); Marloth (1908: 21, 49, 50, 55, IV, V , VI, X , XI); H . C . Taylor (1978: 1-19); Werger et al. (1972: 1-5). Syn.: false macchia (Aco*cks, 1975); macchia (Aco*cks, 1975). The first botanist to use the Afrikaans word fynbos in print appears to have been Bews (1916). It aptly conveys the small leaves and bushy habit of the dominant plants. Most stands of fynbos contain m a n y species, and single-species dominance does not occur other than locally. Taylor (1972) has recorded 121 species of flowering plants from a single 100 m 2 quadrat in a hom*ogeneous stand. Floristics. In most types of fynbos other than extreme variants the following genera and families are usually well represented: Protea, Leucadendron, Leucospermum and Serruria (Proteaceae), Erica, Simocheilus, Philippia and Blaeria (Ericaceae), all genera of Restionaceae, Bruniaceae and Penaeaceae, Aspalathus, Podalyria and Cyclopia (Leguminosae), Phylica (Rhamnaceae), Tetraria, Ficinia and Chrysithrix (Cyperaceae), Diosma and Agathosma (Rutaceae), Cliffortia (Rosaceae), Metalasia, Helichrysum, Stoebe, Elytropappus and m a n y others (Compositae), Lobostemon (Boraginaceae), Polygala and Muraltia (Polygalaceae), Grubbia (Grubbiaceae), and m a n y genera of Liliaceae, Amaryllidaceae and Iridaceae. Physiognomy. Fynbos often contains scattered taller bushes and, less often, widely spaced trees. There is always a conspicuous admixture of monocotyledonous 'switch' plants belonging to Restionaceae, which in some extreme habitats become physiognomically dominant. It
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is the constant presence of Restionaceae, above all else, that distinguishes fynbos from the vegetation of other regions with a broadly similar climate. T h e growth-form of Restionaceae, which is also found in some Cyperaceae, is referred to by Taylor as 'restioid'. They are tufted or rhizomatous plants ranging in height from 20 c m to 2 m or more. Their green, tubular or wiry stems, which bear reduced, non-photosynthetic scale leaves, always persist for more than one year, but usually for less than four. Geophytes are normally plentiful in fynbos but annuals are conspicuous only in the drier types. Grasses are u n c o m m o n today, but according to Aco*cks (1949) were m u c h more abundant before the advent of the European. Except in certain disturbed areas, they n o w contribute little to the appearance of the vegetation. Apart from 'weedy' species, they mostly belong to the 'southern' genera Merxmuellera (Danthonia), Pentaschistis, Ehrharta, Plagiochloa and Lasiochloa. Cyperaceae on the other hand are often abundant. T h e shrubs and bushes of fynbos vary greatly in height and density. They are mostly richly branched and have twisted boles. In typical fynbos true trees are virtually absent. T h e only species with well-defined boles are Leueadendron argenteum, the 'Silver Tree', and Widdringtonia cedarbergensis and W. schwarzii. In a few places on Table Mountain, Leueadendron occurs as an emergent up to 10 m high scattered in typical fynbos shrubland. Under present conditions, Widdringtonia cedarbergensis is normally a small tree 5-7 m high, but can attain a height of 20 m , and formerly frequently did so. It occurs scattered chiefly in rocky places in the Cedarberg between 915 and 1525 m , but never forms closed stands (H. C . Taylor, pers. c o m m . ) . W. schwarzii is confined to rocky ravines in the Baviaanskloof and K o u g a Mountains. It can reach a height of 30 m or more but is usually smaller. Like W. cedarbergensis it does not form a closed canopy. T w o other species, Widdringtonia cupressoides and Olea capensis, which often occur as trees elsewhere, in fynbos are usually branched low d o w n , of bushy habit, and less than 7 m tall.
P. repens, P. susannae, Psoralea pinnata and Wiborgia sericea. Only one arborescent Aloe is a m e m b e r of typical fynbos, namely the 5 m tall, repeatedly branched Aloe plicatilis which occurs in the mountains between French H o e k and the Elandskloof M t s , where the rainfall is about 2000 m m per year. Leaves. T h e foliage of the woody plants is mostly brownish-green or greyish. The leaves are sclerophyllous —small, stiff, thick, coriaceous and entire. They are strongly cuticularized and rich in sclerenchyma with reduced intercellular spaces and often have struts which prevent leaf collapse. T h e lower surface is commonly hairy. T w o leaf-shapes, the ericoid and proteoid, are particularly prevalent. Ericoid leaves are small and narrow with rolled margins. They are found in all types of fynbos and occur in a wide range of unrelated families, e.g. Ericaceae {Erica), Bruniaceae (Bruñid), Polygalaceae (Muraltia), Leguminosae (Aspalathus), Thymelaeaceae (Passerina), Rosaceae (Cliffortia), Rhamnaceae (Phylicá) and Compositae (Metalasia). Proteoid leaves are associated with less branched, usually taller, bushy plants, especially members of the Proteaceae (Leueadendron, Leucospermum, Mimetes, Protea). They are mostly elliptic or oblanceolate, u p to 15 c m long, phyllodic in appearance and very sclerophyllous. Whereas restioids and ericoids are always present in fynbos, proteoids are sometimes absent, especially from drier types and at high altitudes.
Major variants. T h e structural complexity of fynbos decreases with increasing altitude. O n slopes below 900 m , fynbos is dense and usually has three layers: a discontinuous, bushy, proteoid upper layer 1.5-3(4) m tall; an ericoid layer of shrubs up to 1 m , and a ground layer of smaller woody plants, herbs, geophytes, and, especially, Restionaceae. At higher altitudes the fynbos becomes progressively shorter and less stratified and the proteoid element disappears. In the eastern extension of this mountain fynbos, rainfall is more evenly distributed and grasses increase. Dense fynbos thickets up to 5 m tall, dominated by a single species, e.g. Leueadendron salicifolium or Berzelia Bushy species. Only about 50 true fynbos species lanuginosa, sometimes occur on the banks of streams. normally exceed 3.5 m in height, but few of them grow S o m e Restionaceae also become virtually exclusively taller than 6 m , and m a n y are frequently m u c h smaller dominant in hollows and by streams where permanent than this. It is quite exceptional, today, to find them ground water is present. For instance, Chondropetalum occurring at all plentifully as tall plants other than very mucronatum (Dovea mucronata) often forms pure c o m locally. T h e more important a m o n g them are: Cassine parvifolia, Cliffortia arbórea, C. grandifolia, Cryptocarya munities in poorly drained places o n the horizontal angustifolia, Diospyros glabra, Erica caffra, E. cater- sandstone plateau of Table Mountain at about 900 m . Occurrences such as these, however, are unusual. In viflora, E. inconstans, Heeria argéntea, Hyaenanche general fynbos is an exceptionally mixed type. globosa, Laurophyllus capensis, Leueadendron procerum (concinnum), L . eucalyptifolium, L . nobile, L . sabulosum, Leucospermum conocarpodendron, Maytenus oleoides, Fire. Fynbos is very combustible, especially during the Metalasia muricata, Mimetes fimbriifolius, Oldenburgia hottest, driest, windiest time of year, w h e n large areas arbúsculo, Passerina filiformis, Philippia chamissonis, are frequently devastated. It is n o w widely believed that Phylica buxifolia, P. oleifolia, P. paniculata, P. villosa, fynbos evolved in relation to recurrent fires from natural Polygala myrtifolia, Protea arbórea, P. glabra, P. lauri- causes and that fire is necessary for its healthy folia, P. longiflora, P. lorifolia, P. nereifolia, P. obtusifolia, maintenance. Most fynbos species (e.g. Protea arbórea,
The Cape regional centre of endemism
Euclea lancea, most Restionaceae) can sprout again after the fiercest fires. Otherwise their seeds are effectively protected from fire (e.g. Leucadendron salicifolium, Widdringtonia). In the absence of fire m a n y species b e c o m e moribund and die, even in open communities (Restionaceae). S o m e rarities (e.g. Orothamnus zeyheri) have almost become extinct because of over-protection from fire. Aliens. Large areas of fynbos have been invaded and, locally, completely replaced by aliens, originally introduced for reclamation or forestry purposes from other regions with a similar climate. Chief a m o n g them are Hakea acicularis and various wattles (Acacia cyclops, melanoxylon and cyanophylla) introduced from A u s tralia, and Pinus pinaster from the Mediterranean region. Secondary Cape shrubland (Rhenosterbosveld) (mapping unit SO) Refs.: Aco*cks (1975, p. 86-7); Mailoth (1908, p. 98-106); Muir (1929, p. 14-21, 37-49); H . C . Taylor (1978, p. 215-18). Phots.: Aco*cks (1975: 79); Marloth (1908: 21); H . C . Taylor (1978: 18). Syn.: Coastal Renosterveld (Taylor); Coastal Rhenosterbosveld (Aco*cks). There are two main blocks, one in the south, the other in the west. They occur below 300 m between the foot of the mountains and the coastal plain. Rainfall is between 300 and 500 m m per year. T h e soils which are derived from shales are more fertile than those of the mountains and the coast, and have been intensively farmed for centuries. Rhenosterbosveld is usually 1 m tall or less, rarely up to 2 m . Its ability to invade agricultural land has been k n o w n for m o r e than 200 years. In 1775 Sparrman described its encroachment and predicted that it would transform the landscape. Although R h e n o sterbosveld is rich in species, in the southern block the typical fynbos families, Ericaceae, Proteaceae and Restionaceae are lacking; the original vegetation w a s probably evergreen scrub dominated by Olea africana and Sideroxylon inerme with Cussonia spicata, Diospyros dichrophylla, Pterocelastrus tricuspidatus etc. T h e southern Rhenosterbosveld is m u c h m o r e grassy than the western and m a n y of its grass species, such as Hyparrhenia hirta, are widespread in the tropics. T h e western block has a greater admixture of fynbos species and the characteristic grasses, such as Lasiochloa echinata and Pentaschistis patula, are non-tropical annuals. Bushy species are fewer but include Olea africana. Coastal bushland and thicket (mapping unit 50) O n the south coast certain scrubs and trees, which are not true fynbos species, locally form thicket or scrub
135
forest up to 10 m tall. They include: Cassine peragua, *Euclea racemosa, E. tomentosa, E. undulata, Maytenus heterophylla, Myrsine africana, *Olea africana, Chionanthus foveolatus, Pterocelastrus tricuspidatus, Putterlickia pyracantha, Rhus crenata, R. glauca, R. laevigata, *R. lucida, *R. tomentosa, *Sideroxylon inerme, Tarchonanthus camphoratus and Zygophyllum morgsana. They are often associated with big restioids, especially Willdenowia stricta, and tall fynbos species of karroid appearance, e.g. Eriocephalus racemosus. Tall scrub (3-6 m high) in H e r m a n u s District has been described by Taylor (1961). It is partly serai to forest of Afromontane affinity but is also an edaphic subclimax o n the dry northern slopes of limestone outcrops where the soil is shallow and well drained. It is m a d e up of those species marked with an asterisk in the list above together with Carissa bispinosa, Chrysanthemoides monilifera, Chionanthus (Linociera) foveolatus, Maytenus heterophylla and Osyris sp. In the western coastal fynbos, bushy species of tropical affinity are fewer than in the southern belt, principally Diospyros austro-africana subsp. rugosa, Euclea natalensis subsp. capensis, E. racemosa, Maytenus heterophylla, Osyris sp., Pterocelastrus tricuspidatus, Putterlickia pyracantha, Rhus glauca and R. mucronata. They occur as scattered individuals 2 - 3 m high or locally form small thickets. T h e m o r e conspicuous true fynbos species are Leucadendron salignum, Metalasia muricata, Protea repens, Thamnochortus erectus, T. spicigerus and Willdenowia striata.
Riparian bushland and thicket (mapping unit 50) T h e lower and less steep watercourses are fringed with 5-7 m tall, dense thickets of Brabeium stellatifolium, Freylinia oppositifolia and Metrosideros angustifolia. At higher altitudes Rapanea melanophloeos, Kiggelaria africana, Maytenus acuminata, Olea africana, Olinia and Podocarpus elongatus are the characteristic species. Cunonia capensis, Hartogia capensis, Ilex mitis and Maytenus oleoides are c o m m o n to both types.
T h e transition to Karoo (mapping unit 50) Along the inner margin of the C a p e Region there is a narrow band of Arid Fynbos which forms the transition from typical C a p e to typical Karoo vegetation. Ericaceae are absent and Proteaceae and Restionaceae though conspicuous, especially the latter, are few in species. Typical shrubby Karoo genera such as Chrysocoma, Hermannia, Euryops, Pteronia, Eriocephalus, Selago, Walafrida, and Lightfootia are well represented. Succulents, including Euphorbia mauritanica and Aloe ferox (in the east), are often present.
VI
The Karoo-Namib regional centre of endemism
Geographical position and area
Geographical position and area
Geology and physiography
This region occupies the central, northern and northwestern parts of the C a p e Province immediately to the north of the C a p e floristic Region (but also has important exclaves within the latter), mostly north of 33° S. and between 17° and 25° E . It extends northwards as an increasingly narrow band along the entire length of Namibia into south-west Angola to about 11 ° S. This northern extension of the K a r o o - N a m i b Region not only includes the coastal plain but also the escarpment of the interior plateau and locally the fringes of the plateau itself. (Area: 661000 k m 2 . )
Climate Flora Mapping units Bushy Karoo shrubland Succulent Karoo shrubland Dwarf Karoo shrubland Montane grassy Karoo shrubland Vegetation Semi-desert vegetation of the Karoo Karoo shrubland Dwarf succulents and succulent shrubs Arborescent succulents Non-succulent bushes, bushy trees and tall shrubs Dwarf non-succulent shrubs Grasses Geophytes and annuals Karooriparianscrub forest The transition to Tongaland-Pondoland evergreen bushland The Namib desert The Outer Namib fog desert Sand dunes Gravel desert Rocky outcrops The Inner Namib desert The Welwitschia bainesii transition zone River-bed communities The desert of Mossamedes
Geology and physiography Both the geology and the physiography are very diverse. Altitude ranges from sea-level to 2695 m . T h e region includes four of the Geomorphic Provinces of King (1951), namely C a p e Middle Veld, Karoo, N a m i b , and Kaokoveld, as well as parts of 'Highveld' and Damaraland. In the interior of the C a p e Province the surface is formed of the Karoo System and is extremely even except where broken by dolerite dykes, sills and other intrusions. T h e soils, mostly derived from D w y k a tillite and dolerite, are clayey and tend to accumulate salts. Brackish seasonal s w a m p s or 'vloere' are extensive. In the North-West C a p e the Karoo beds have been removed to expose granite and other primitive rocks with numerous later igneous intrusions, which provide abundant sand for distribution by wind. This area is mostly of subdued relief but rugged mountains occur in western N a m a q u a l a n d and in the gorge tract of the Orange River. T h e N a m i b desert occupies a coastal peneplain, extensive areas of which are covered with moving sand of recent origin. Elsewhere, granite, gneiss, or Stormberg lava outcrop at the surface. Further inland in Namibia the rocks are very varied and give rise to a variable relief.
Climate Rainfall in the N a m i b Desert is less than 100 m m per year. Elsewhere it rarely exceeds 250 m m . Seasonality of
The Karoo-Namib regional centre ofendemism
rainfall varies greatly. West of a line running from Spencer Bay through Calvinia to Sutherland more than 60 per cent of the rain falls in winter. East of a Une from S w a k o p m u n d through Pofadder and Fraserburg to Willimore more than 60 per cent falls in s u m m e r . Nevertheless, in most of the ' s u m m e r rainfall' parts of the Region there is more dry-season precipitation than in most parts of the Zambezian Region, or, as in the coastal belt, dry-season mists are frequent (see page 141). There is considerable variation in the amount and distribution of rainfall from year to year, especially in the driest parts. Even in the wetter parts of the s u m m e r rainfall belt, winter influences are dominant about one year in twelve. T h e coastal belt is frost-free except in the south, where occasional light frosts occur in July as at Port Nolloth. Further inland in southern Namibia and throughout the interior of the Cape the frost period lasts for 5-6 months, though the m e a n m i n i m u m temperature of n o m o n t h is below zero except in the extreme east towards the contact with the climatic limit of Highveld grassveld. (See Fig. 12.)
Flora There are about 3500 species, of which more than half are endemic. Endemic family. Welwitschiaceae (1 species, Welwitschia bainesii). Other characteristic families. Asclepiadaceae: Stapelieae (6 endemic genera and c. 160 endemic species). Aizoaceae (Mesembryanthemaceae) (95 endemic genera and c. 1500 endemic species).
137
Aloe speciosa, Carissa haematocarpa, Crassula portulacea, Euclea undulata, Euphorbia grandidens, Montinia caryophyllacea, Portulacaria afra, Schotia afra and S. latifolia. Tree species which extend into the K a r o o - N a m i b Region from the Zambezian Region, and in some cases from m u c h further north, include Acacia mellifera subsp. detinens, A. erioloba, A. karroo, Boscia albitrunca, Diospyros lycioides, Dodonaea viscosa, Euclea crispa, Pappea capensis and Ziziphus mucronata. Grasses which extend at least as far as the Z a m bezian Region include Cymbopogon plurinodis (pospischilii), Eustachys paspaloides, Fingerhuthia africana, Hyparrhenia hirta, Schmidtia pappophoroides and Themeda triandra.
Mapping units 51. Bushy K a r o o shrubland. 52. Succulent Karoo shrubland. 53. D w a r f Karoo shrubland. 57a. M o n t a n e Grassy Karoo shrubland. 74. T h e N a m i b desert (see below). T h e four mapping units into which the Karoo has been divided have been adapted from those of Aco*cks (1975). There is a fair amount of floristic information, but for large areas ecological information is virtually nonexistent. For this reason, after the mapping units have been briefly characterized, the Karoo, apart from its riparian and transitional vegetation, is treated as a single continuum. Bushy Karoo shrubland (mapping unit 51)
Endemic genera (in addition to above). About 60 including Adenolobus (2 species), Arthraerua (1), Augea (1), Ceraria (5), Didelta (2), Grielum (6), Kaokochloa (1), Leucosphaera (2), Monelytrum (1), Nymania (1), Phaeoptilum (1), Phymaspermum (9), Sisyndite (1), Xerocladia (1).
Refs.: Aco*cks (1975, p. 59-63, 71-5); Barbosa (1970, p. 24551); Giess (1971, p. 9-12); de Matos & de Sousa (1970); White (MS, 1973). Phots.: Aco*cks (1975: 54, 55, 64-7); Cannon (1924: 8b & c, lia, 19b, 21b); Giess (1971: 21, 22, 24-7); Marloth (1908: 105, 107, 108, X V I , XVIII); de Matos & de Sousa (1970: 4); Shantz& Turner (1958: 11,12,13,15). Syn.: karroid broken veld; Namaqualand broken veld; Orange River broken veld (all of Aco*cks, 1975).
Endemic species. T h e following genera have important concentrations of endemic species: Aloe, Anacampseros, Babiana, Chrysocoma, Cotyledon, Crassula, Eriocephalus, Euphorbia, Gasteria, Haworthia, Hermannia, Pentzia, Pteronia, Sarcocaulon, Stipagrostis, Tetragonia, Zygophyllum. S o m e of these genera, e.g. Pteronia, are almost confined to the K a r o o - N a m i b Region. A t the other extreme, some, e.g. Euphorbia, are cosmopolitan.
This is shrubland dotted with small bushy trees and large shrubs. It occurs in the Great Karoo, the Little Karoo, Robertson Karoo, and o n the rocky hills of Namaqualand and the lower valley of the Orange River and northward to south-west Angola. Succulents are usually abundant, especially in the south. Non-succulent dwarf shrubs are always present but are usually subordinate to the succulents. Grasses though inconspicuous are represented by m a n y species.
Linking elements. A t the species level there is little interminghng with the Cape flora. Most linking species extend to the east or the north or both. Species c o m m o n to the K a r o o - N a m i b and Tongaland-Pondoland Regions, and in some cases extending slightly beyond their combined area, include
Succulent Karoo shrubland (mapping unit 52) Refs.: Aco*cks (1975, p. 69-71). Phots.: Aco*cks (1975: 62); White, Dyer & Sloane (1941: 190d, 276).
138
Vegetation of the floristic regions
Ç?*i* r-f _«_c-e-c.-G-e-*—^.e-o-o-e-
FIG. 12. Climate and topography of the Karoo-Namib regional centre of endemism (VI) A . North of Cunene River. B . South of Cunene River
The Karoo-Namib regional centre ofendemism
Succulent Karoo shrubland, except for secondary types, is largely confined to the sandy coastal plain of Namaqualand and the heavier stony soils of the foothills of the Namaqualand escarpment. There is an important outlier, enclosed by high mountains in the rain shadow valleys of the Tanqua and D o o m Rivers. These areas occur mostly below 610 m and are frost-free. The annual rainfall is less than 200 m m and falls in winter. In the coastal areas the effects of aridity are reduced by sea mists. Succulents are dominant throughout. They range in height from almost subterranean species to shrubs 2 m or more tall, but the average height is between 0.3 and 1 m . In the driest and most degraded areas ground cover is very low but elsewhere m a y be as high as 50 per cent. Large shrubs and bushes are virtually absent except along therivers.Dwarf non-succulent shrubs are represented by numerous species but they contribute little to the physiognomy. Grasses are few in species and usually inconspicuous.
139
Vegetation The desert vegetation of the N a m i b , and the semi-desert vegetation of the Karoo, including its northern extension into Angola, are described separately below. A richly illustrated review of the literature on the Karoo-Namib Region has been published by Werger (Werger, 1978ft, in Werger (éd.), 1978c). Semi-desert vegetation of the Karoo Except along the larger watercourses, which support a fringe of scrub forest, bushland, or thicket, almost the whole area is covered with shrubland. Only locally in the transition to wetter regions is the vegetation sufficiently luxuriant to be classified as bushland. Although Karoo shrubland includes a wide range of physiognomic types there is insufficient published information to justify their separate description in a general work such as this. Karoo shrubland
Dwarf Karoo shrubland (mapping unit 53) Refs.: Aco*cks (1975, p. 63-9, 76-7); Adamson (1938a, p. 17988); Marloth (1908, p. 280-90). Phots.: Aco*cks (1975: 56-61), Adamson (1938a: 14); Marloth (1908: 17a); Shantz& Turner (1958: 19&20). Syn.: Arid Karoo; False Arid Karoo; Central Lower Karoo; Central Upper Karoo; Western Mountain Karoo (all of Aco*cks, 1975). This is the most extensive type of Karoo shrubland. It is dominated by dwarf shrubs, most of which belong to Compositae. It occupies the Arid and Central Karoo and the lower slopes of the Karoo mountains. Bushes and trees are absent. Large shrubs are few in species and local, and are represented chiefly by Rhigozum trichotomum. Succulents are always present but are mostly inconspicuous; there are relatively few species. Grasses are more abundant than in other types and increase towards the east. T h e soils are often slightly saline, and halophytes, particularly Salsola tuberculata, are widespread, and locally dominant. In the Arid Karoo there are enormous brackish flats or 'vloere', in some jcases covered with Salsola aphylla and other halophytes, but elsewhere practically bare. Montane grassy Karoo shrubland (mapping unit 57a) Published information is almost non-existent. Aco*cks (1975, p . 81) suggests that the original vegetation was formerly more grassy. Today Merxmuellera disticha and M . stricta are the chief relict grasses; Karoo shrublets belonging to Chrysocoma, Eriocephalus, Pentzia, Ruschia etc., as well as Elytropappus rhinocerotis, are abundant.
Except very locally the shrubs are less than, and frequently m u c h less than, 2 m tall. Over extensive areas, however, the landscape is dotted with larger woody plants, either arborescent succulents, or non-succulent bushes or bushy trees. These taller plants rarely exceed 5 m in height. They are chiefly confined to places, usually rocky, where the water supply is increased by run-off from surrounding slopes. Hence they are not uniformly distributed and the landscape is a mosaic of bushed and unbushed areas. In the former the bushes vary from 5 to 100 m apart. They reach their greatest luxuriance in places where there is mist condensation, such as in the Richtersveld. In Karoo shrubland there is as wide a diversity of growth forms as in Cape fynbos, but the subject has been little studied. For convenience, in addition to grasses and forbs, the following principal types m a y be recognized: (a) dwarf succulents and succulent shrubs; (b) arborescent succulents; (c) dwarf non-succulent shrubs; (d) taller non-succulent shrubs, bushes and bushy trees. Their relative abundance varies greatly. Dwarf succulents and succulent and non-succulent shrubs occur throughout. Arborescent succulents and non-succulent bushes and bushy trees, however, are more or less confined to 'broken veld' (see mapping unit 51 above). Fewer bushy species occur in the southern broken veld of the Little and Great Karoo than elsewhere. Dwarf succulents and succulent shrubs Stem succulents are represented by species of Euphorbia, Asclepiadaceae (especially Stapelia, Caralluma, Hoodia, Huernia and Trichocaulon), Senecio {'Kleinia'), and by species of Sarcocaulon and Pelargonium which shed their small mesomorphic leaves during drought. Most species are less than 1 m tall, but the largest Euphorbias are up
140
Vegetation of the floristic regions
to 2 m . The cactoid E. avasmontana is conspicuous in the mountainous country on both sides of the Orange River. O f the non-spinous species E. mauritanica is the most widespread. E. gummifera and E. gregaria are abundant in parts of Namaqualand and Namibia. Leaf-succulents are pre-eminently represented by Mesembryanthemaceae. T h e most prevalent are small shrubs from 30 c m to 1 (2) m tall, especially species of Ruschia, but the annuals and stone plants (Lithops, Titanopsis, Argyroderma etc.) are locally important. Other widespread genera with m a n y succulent species include Anacampseros (Portulacaceae), Cotyledon and Crassula (Crassulaceae), Aloe, Haworthia and Gasteria (Liliaceae), Lycium (Solanaceae) and Zygophyllum (Zygophyllaceae).
africana and Rhigozum trichotomum occur in N a m a qualand and the Orange River Valley. S o m e , e.g. Acacia redacta, are endemic to this region. Others such as Parkinsonia extend as far north as Angola. Species confined to the northern part of the K a r o o N a m i b Region include Acacia montis-usti, A. robynsiana, Adenolobus pechuelii, Euphorbia guerichiana, Rhigozum virgatum and several species of Commiphora. Most of the above are from 2.5 to 4 m tall and are branched from the base or have short irregular boles. Acacia erioloba, however, is sometimes taller and has a straight bole. Dwarf non-succulent shrubs
South of the Orange River nearly all of the most abundant dwarf shrubs belong to non-tropical genera, most of which are virtually confined to South Africa or have Aloe dichotoma (up to 5 m high), which extends from their main or subsidiary centres of endemism there. Namaqualand almost as far north as Windhoek, is the They include Aster, Berkheya, Chrysocoma, Didelta, most abundant species. The more sparsely branched A. Eriocephalus, Euryops, Garuleum, Helichrysum, Lightpillansii (10 m ) is almost confined to the Richtersveld. footia, Osteospermum, Pentzia and Pteronia in C o m Cotyledon paniculata (3 m ) , Cerararia namaquensis (5 m ) , positae, and Galenia, Hermannia, Lebeckia, Nestlera, and the unbranched Pachypodium namaquanum (5 m ) Plinthus, Selago, Sutera, Wahlenbergia and Walafrida in extend from Namaqualand into southern Namibia. In other families. T h e relative importance of these genera northern Namibia the last two are replaced by C . steadily diminishes towards the tropics and only a few longepedunculata and Pachypodium lealii respectively. species, e.g. Pteronia glauca, extend into northern Towards the northern end of the K a r o o - N a m i b Region Namibia. In the northern part of the K a r o o - N a m i b two succulent Euphorbias, E. currorii (10 m ) and E. Region small shrubs and suffrutices (and also herbs) eduardoi occur. There are also several species with belonging to tropical genera such as Barleria, Blepharis, enlarged water-storing stems, including Cyphostemma Crotalaria, Hibiscus, Indigofera, Monechma, Petalidium, currorii (7 m ) , Moringa ovalifolia (7 m ) and Sesamo- Pterodiscus, Ruellia and Tephrosia become more thamnus guerichii and S. benguellensis (5 m ) . South of important. Namaqualand and the Orange River Valley arborescent T h e dwarf non-succulent Karoo shrubs are rarely succulents are m u c h less plentiful and are represented more than 1 m tall and commonly are no more than only by Crassula arborescens (3 m , Little Karoo 25 c m . In heavily grazed areas they m a y be no more eastwards) and Portulacaria afra (4 m , Little Karoo than 10 c m . eastwards). Arborescent succulents
Grasses Non-succulent bushes, bushy trees and tall shrubs
About 130 species of grass occur in the K a r o o - N a m i b Region, and 80 or so are confined or almost confined to it. T h e endemic species belong to 29 genera, of which There are about 100 species in this group. Dodonaea Aristida and Stipagrostis are the most important. viscosa, Euclea undulata, Nymania capensis, Pappea Grasses occur throughout the Karoo but are only locally capensis, Rhigozum obovatum, Rhus undulata and Schotia physiognomically dominant. afra are widespread south of the Orange River and extend at least a short w a y further north into Namibia. There can be little doubt that the importance of Ehretia rígida, Boscia albitrunca, B. foetida, Acacia grasses in relation to woody plants has declined within erioloba and A. mellifera subsp. detinens and Grewia recent times owing to overgrazing, and there is some photographic evidence in support of this (Shantz flava, which are widespread in the drier parts of South & Turner, 1958), but it is extremely unlikely that pure tropical Africa, extend south to beyond the Orange grassland anywhere represents the climax vegetation, River. In northern Namibia and south-west Angola except possibly very locally on deep sandy soils and in a m a n y more Zambezian linking species enter the K a r o o few other edaphically favourable places. N a m i b Region and some, e.g. Colophospermum mopane, almost reach the sea. Grasses are m u c h less conspicuous in succulent Acacia redacta, Adenolobus garipensis, Commiphora Karoo and bushy Karoo than in dwarf non-succulent capensis, C. gracilifrondosa, C. namaensis, C. oblanKaroo. In the latter the most conspicuous species are ceolata, Diospyros ramulosa, Ficus cordata, F. guethe silvery-white desert grasses, so called because of richiana, Heeria concolor, H . crassinervia, Parkinsonia the bleached appearance of their dead persistent
The Karoo-Namib regional centre ofendemism
141
inflorescences. T h e principal species are Aristida diffusa, page 201), occur in the Southern and Eastern Cape on Eragrostis lehmanniana, Stipagrostis brevifolia, S. ciliata, steep mountain slopes where annual rainfall is 2 5 0 S. namaquensis, S. obtusa and S. uniplumis. Normally 300 m m . They are intermediate in structure and floristic they are perennial, but they sometimes complete their composition between the drier types of Tongalandgrowth-cycle within a single season. In times of drought Pondoland evergreen bushland and the more luxuriant Stipagrostis brevifolia, which is the hardiest of all the types of bushy K a r o o - N a m i b shrubland. T h e Noorsveld plants of the Arid Karoo, sheds its leaves and is of the Eastern Cape (Aco*cks, 1975, p . 58; V a n der Walt, transformed into a small w o o d y shrublet. After good 1968), which occurs at lower altitudes and has a slightly rains grasses sometimes temporarily overtop and conceal lower rainfall, is similarly transitional. the shrublets they are growing with. Elsewhere, grasses m a y besóme physiognomically dominant following The N a m i b desert extreme overgrazing because they regenerate more (mapping unit 74) rapidly from seed than their associates. Refs.: Giess (1968a, 1971); Marloth (1909); Walter (1971, p. 338-74); Werger (19786). Geophytes and annuals Phots.: Adamson (1938: 16); Coetzee & Werger (1975: 13-17); Giess (1968a: 1-6); Marloth (1909: 4); Walter (1971: 209, Geophytes and annuals are well represented in the 225); Werger (19786: 3, 4, 6,10,11). Karoo flora, especially in the winter-rainfall area of Namaqualand. They are conspicuous only after periods T h e N a m i b desert runs the whole length of Namibia and of good rainfall when, for a few weeks, their attractive continues a short distance further north into Angola as flowers transform the appearance of the veld. Important the desert of Mossamedes, and a short distance to the genera of geophytes include Babiana, Bulbine, Hornería, south along the coast of Namaqualand. For most of its Lachenalia, Lapeirousia and Oxalis. The most abundant length it is about 100 k m wide and extends from the annuals belong to species of Arctotis, Cotula, Dimorcoastline to the foot of the scarp which delimits the photheca, Felicia, Osteospermum, Senecio, Ursinia, Veniinterior highlands of southern Africa. The boundary of dium (all Compositae) and Heliophila, Hermannia and the N a m i b is arbitrarily defined by the 100 m m isohyet, Grielum. but the greater part of it receives very m u c h less rainfall In overgrazed areas, especially on sandy soils than this. towards the northern limits of the Karoo, the noxious Giess (1971) divides the N a m i b into three. T h e weed Tribulus zeyheri forms similar extensive colourful Northern N a m i b extends as far south as the H u a b River. carpets. T h e Central N a m i b lies between the H u a b and Kuiseb Karoo riparian scrub forest Acacia karroo is gregarious and widespread in the Karoo and is often the only tree present, especially in the interior. Nearer the Atlantic coast, as in the PofadderAugrabies sector of the Orange River, the riparian bushland flora is more diversified, and Acacia karroo is sometimes rarer there than Pappea capensis (6 m ) , Euclea pseudebenus (8 m ) , Tamarix usneoides (7 m ) , Diospyros lycioides (7 m ) , Rhus undulata (3 m ) and Euclea undulata (6 m ) . Other associates include Diospyros aco*cksii, Rhus lancea, Combretum erythrophyllum and Ziziphus mucronata. Towards the northern end of the K a r o o - N a m i b Region several species which are widespread in the Zambezian Region (and sometimes also elsewhere) as quite large trees penetrate far into the desert along watercourses, usually in the form of bushy trees. They include Acacia albida, which is sometimes kept small by browsing zebra, A. erioloba, Colophospermum mopane, Combretum apiculatum, C. imberbe, Ficus sycomorus, Sterculia africana and Ziziphus mucronata. The transition to Tongaland—Pondoland evergreen bushland Communities dominated by Portulacaria afra (Aco*cks, 1975, p . 58-9), and k n o w n as 'Spekboomveld' (see
Rivers. T h e Southern N a m i b , which extends into northern Namaqualand, experiences s u m m e r rainfall in the northern part and winter rainfall in the south. Walter (1971) distinguishes between the Outer N a m i b , which receives frequent fogs, and the fog-free Inner Namib. T h e drier parts of the N a m i b receive rainfall only rarely, but the outer coastal belt is characterized by m a n y days of fog. In the southern part as far north as Luderitz Bay such rain as there is falls in winter; further north it falls in summer. In the coastal part of the Central N a m i b near S w a k o p m u n d the m e a n annual rainfall is only 10 m m , and on average rain occurs only once in two years, but at remote intervals high rainfall is experienced. In 1934, for instance, there was a rainfall of nearly 150 m m ; by contrast in certain years rainfall is hardly measurable. A t S w a k o p m u n d there are 94 to 215 fog days per a n n u m and the frequency is no less in the Southern N a m i b . Their effects rarely penetrate as far as 50 k m inland. T h e fog belt which is carried constantly above the cold offshore Benguela current is brought inland by south-westerly winds during the night and only disperses during the day w h e n the desert soil becomes heated. T h e precipitation from individual fogs is mostly less than 0.1 m m and is never more than 0.7 m m . The total annual precipitation from fog is between 40 and 50 m m but in most places this is of no significance to vascular
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Vegetation of the floristic regions
plants since the water only penetrates to 3.5 c m a n d evaporates immediately the fog dissipates. T h e situation is different, however, where fog is condensed o n rock faces. T h e n water from a large catchment area runs into crevices a n d vascular plants can b e c o m e established, even though the true rainfall is too low to measure. T h e plants g r o w mainly during the cool season, w h e n fogs are frequent, a n d not during the s u m m e r w h e n there m a y b e occasional falls of rain. T h e fog-water contains s o d i u m chloride a n d the soils of the Outer N a m i b are brackish as far as the inland limit of coastal fog, while the Inner N a m i b has n o saline soils. The Outer Namib fog desert Sand dunes Phots.: Giess (1971: 1, 2,14, 15). There are t w o m a i n areas, o n e in the Northern N a m i b a n d the other in the northern half of the Southern Namib. The former is up to 40 k m wide and extends to the north of the Cunene River far into Angola. The extremely sparse vegetation consists of isolated plants of Barleria solitaria, Ectadium virgatum, Indigo/era cunenensis, Merremia multisecta, Petalidium angustitubum, P . giessii, a n d the grasses Stipagrostis ramulosa a n d Eragrostis cyperoides. South of S w a k o p m u n d a n d extending as far south as Luderitzbucht there is a large belt of shifting dunes about 320 k m long a n d 120 k m wide. Although botanically unexplored it is believed to b e virtually without vegetation. T h e few species k n o w n to occur include Monsonia ignorata, Trianthema hereroensis, Stipagrostis gonatostachys a n d S. sabulicola. In the Central N a m i b there is frequently a narrow strip of small dunes u p to 2 0 0 m wide with scattered cushions of Psilocaulon salicornioides, Salsola aphylla a n d S. nollothensis. Gravel desert Phots.: Giess (1968a: 1; 1971: 7,16); Marloth (1909: 4). N o r t h of S w a k o p m u n d gravel desert occupies m o s t of the outer parts of the Central N a m i b . Fifty per cent of the surface is covered with a stony pavement. T h e soil is cemented into a rock-hard layer b y deposition of lime a n d g y p s u m forming a hardpan at a depth of 1-5 c m . Soil wetting occurs to this depth. All the stones are covered with colourful folióse and crustose lichens, such as species of Parmelia a n d Usnea a n d Teloschistes capensis. Otherwise the gravel desert is normally devoid of vegetation. After the heavy rains of 1934 n u m e r o u s plants appeared. Their m e a n cover w a s 2 0 per cent and, in small depressions, as m u c h as 5 0 - 9 0 per cent. Typical plants were Psilocaulon (Mesembryanthemum) salicornioides, Mesembryanthemum cryptanthum (Hydrodea bossiana), Drosanthemum luederitzii (paxianum),
Aizoanthemum (Aizoori) dinteri a n d Zygophyllum simplex. These are all ephemeral halophilous succulents with extremely l o w rates of transpiration. T h e exceptionally high rainfall of 1934 enabled s o m e of t h e m to remain alive for m o r e than a year but eventually all s u c c u m b e d to drought. A few individuals of t w o perennial species, Arthraerua leubnitziae a n d Zygophyllum stapfii, which are normally confined to drainage lines, colonized the plains but did not persist. A n n u a l grasses, including Stipagrostis hermannii, S. namibensis a n d S. subacaulis, also appear plentifully after adequate rain has fallen. T h e Southern N a m i b has a higher rainfall than the Central N a m i b , a n d perennial species such as the stem succulent, Euphorbia gummifera, a n d small shrubs of Zygophyllum retrofractum a n d Sarcocaulon spinosum occur in places o n gravelly flats. Rocky outcrops In the outer part of the Central N a m i b , apart from ground-water areas, rocks provide the only habitats where perennials can survive. T h e rocks are usually covered with lichens. T h e following dwarf succulents predominate a m o n g the vascular plants rooted in crevices: S t e m succulents: Trichocaulon clavatum (dinteri), T. pedicellatum, Hoodia currorii. Leaf succulents: Lithops spp., Anacampseros Aloe asperifolia, Cotyledon orbiculata.
albissima,
S t e m succulents with deciduous leaves: Pelargonium otaviense (rossingense), Sarcocaulon mossamedense (marlothii), Othonna protecta, Senecio longiflorus, Adenia pechuelii. The Inner Namib desert Phots.: Giess (1971: 5, 8); Walter (1971: 209). Information o n the region is sparse because most accounts of the N a m i b desert are generalized and d o not d r a w a clear distinction between the Outer a n d Inner N a m i b or between the latter a n d adjoining semi-desert regions. According to Walter (1971) the plains of the Inner N a m i b are devoid of halophytes, a n d in thenplace occur grasses, chiefly species of Stipagrostis, associated with herbaceous Compositae a n d Acanthaceae. Succulents are represented b y a single species, Sesuvium sesuvioides (digynum) (Aizoaceae). Stipagrostis obtusa is probably the m o s t generally occurring dominant. It is often m i x e d with S. ciliata in the sandier parts a n d with Eragrostis nindensis o n m o r e stony or gravelly soils. Kaokochloa nigrirostis sometimes forms pure stands in the Northern N a m i b . South of the O r a n g e River similar desert grassland occurs o n Kalahari S a n d between Okiep a n d Pofadder. T h e rainfall is less than 4 0 m m . Stipagrostis brevifolia is the dominant species. It is a low shrubby plant with the
The Karoo-Namib regional centre ofendemism
perennating shoots branched above ground. During the dry season it sheds its leaves. At this time, when no inflorescences axe present, the plants have the appearance more of dicotyledonous shrubs than of typical grasses. Although the tufts are widely spaced (mean distance c. 1 m ) and rather small, being only 20-30 c m high and wide, the landscape from a distance looks as if it is densely shrubby. There are few associated species. The large shrub Parkinsonia africana m a y occur as widely scattered individuals. Typical Karoo dwarf shrubs are virtually absent, being represented by a few individuals of Lycium and Hermannia. After a good fall of rain, annuals, especially annual grasses and Tribulus zeyheri, are plentiful. The Welwitschia bainesii transition zone Refs.: Bornman et al. (1972-73); Giess (1969); Walter (1936; 1971, p. 369-73). Phots.: Bornman et al. (1972: 2-3); Giess (1969: 12-18; 1971: 6,11). The distribution of Welwitschia bainesii, one of the most remarkable plants in the world, has been described in detail by Kers (1967), Giess (1969), and Barbosa (1970). It extends from the Kuiseb River just south of the Tropic of Capricorn to San Nicolau (14°20'S.) in southern Angola. In contrast to earlier beliefs its range is n o w k n o w n to be almost continuous. It has been studied most in the Central N a m i b towards the southern limit of its range where it is the characteristic species of the transition zone between the Outer and Inner N a m i b , and occupies a narrow strip about 50 k m from the coast between the Kuiseb and Swakop Rivers (Walter, 1971; Bornman et al, 197273). According to Walter, fog is rare here (but see below) and is of little significance to the plant, although light summer rains are frequent. In the Kuiseb-Swakop area the distribution of Welwitschia is not related to ground water and its taproots only go d o w n to a depth of 1-1.5 m . It does not occur on the plains themselves, which are covered, after rains, with annual grasses such as Stipagrostis subacaulis and annual forms of perennial species such as S. hochstetterana. Welwitschia is found in broad flat channels in the plains which are so shallow as to be barely discernible. These channels receive floodsheet waters from higher areas, and the soil becomes moist to a depth of 1.5 m . This subsurface moisture can be retained for years. T h e Welwitschia plants are usually more than 20 m apart and associated plants other than ephemerals are sparse. Welwitschia can store a certain amount of water but has no specialized water-storing tissue. With lack of water the leaves die except for their meristematic bases. Welwitschia also occurs on slopes covered with coarse scree and in crevices in weathered rock but the plants are isolated and not very vigorous (Walter, 1971). The ecology of Welwitschia has been studied by Bornman and his collaborators (Bornman et al., 1972-
143
73), both in the field and in the laboratory. Where m e a n annual rainfall is only 25 m m , precipitation from coastal fog is equivalent to a further 50 m m . Bornman has suggested that, contrary to the statements of Walter, some of this fog condensation is absorbed by the leaves, probably through the stomata, but the evidence is inconclusive (L. Leyton, pers. c o m m . ) . The plant body of Welwitschia resembles a large fibrous carrot. T h e stem is up to 1.5 m tall and up to 10.8 m in circumference. Bornman estimates that the largest individuals are 2500 years old. There are only two leaves, which grow from a terminal groove in the photosynthetic tissue of the stem. They persist throughout the entire life of the plant and under favourable conditions grow from the basal meristems at the rate of 13.8 c m per year, so that the oldest plants would be capable of producing leaves well over 100 m long if it were not for their intermittent growth and continuous dieback from the extremities. W h e n the leaf tip comes into contact with the ground it withers following the death of protoplasts caused by the high surface temperatures, and it becomes frayed from being scoured on the gravel surface by winds. However, even under gale force conditions, the living part of the leaf, which is rarely more than 3 m long, remains remarkablyrigidand stable. Welwitschia frequently forms groups of individuals of equal size, which are presumably of similar age. Bornman suggests that germination follows a freak downpour or series of downpours of approximately 25 m m . Germination will not take place until an inhibitor has been leached from the seed. The equivalent of 6.25 m m of rain is needed for this. Walter (1936, 1971) implies that the transition zone between the Outer and Inner N a m i b is the most characteristic habitat of Welwitschia, but this is n o w k n o w n to be true only for the southernmost localities and some of the northern ones. It is not valid for the general distribution of the species (Kers, 1967). Welwitschia can occur within 8 k m of the coast and extends up to 144 k m inland. Its altitudinal range is from 100 to 900 m and it tolerates a wide range of precipitation and soil salinity and grows in a variety of vegetation types. Welwitschia is not confined to the N a m i b Mossamedes desert nor to the K a r o o - N a m i b Region, but extends some way into the Karoo-Namib/Zambezia transition zone. At its eastern limit in Namibia near the small town of Welwitschia, where the rainfall is 200 m m per year, it occurs on gravel soils of deltaic deposits. Here, it is most abundant in the shade ofriparianbushland dominated by Colophospermum mopane and Terminalia prunioides. Individuals are relatively large but are m u c h smaller than the grotesque giants of the desert. O n the stony flats between the watercourses, Welwitschia is the only 'woody' plant. T h e associated vegetation is composed of annual grasses, principally Stipagrostis hirtigluma and Anthephora schinzii, which sometimes completely conceal it. Welwitschia also occurs in this area on rocky sandstone hills covered with a relatively
144
Vegetation of the floristic regions
dense bushland dominated by Colophospermum mopane, Terminaliaprunioides, Acacia spp. and Commiphora spp. At its eastern limits in Namibia Welwitschia is confined to compact gravel soils or rocky places. It cannot establish itself o n less compact soils, since young plants are uprooted and washed away by run-off from violent rainfall (Kers, 1967). River-bed communities Phots.: Giess(1971: 3, 9,10,11,12); Walter (1971: 225).
Welwitschia is absent from the coastal belt of mobile dunes south of Porto Alexandre, but otherwise is scattered throughout. T h e vegetation of the Reserve Parcial de Mossamedes towards the northern limit of the desert has been described by de Matos and de Sousa (1970), w h o recognize the following main types: 1. Near the coast on saline soils there are halophytic communities characterized by Salsola zeyheri, Sesuvium spp., Suaeda fruticosa, Scirpus littoralis and Asthenatherum {Danthonia) forskalii.
A s in all deserts, erosion channels and dry valleys are the 2. A little further inland, o n mesa-like terraces and in most favourable habitats for plants. W h e n the rivers ravines o n less saline, calcareous and gypsaceous soils, flow, the sand in their otherwise dry channels becomes the following are found: Aizoon virgatum, A. mossawetted to an appreciable depth and then remains d a m p medense, Euphorbia bellica, Zygophyllum orbiculatum, Z . for several years. Most river beds are at least slightly simplex, Rhynchosia candida, Indigofera daleoides, Geisaline but the degree of salinity is variable and depends geria spinosa and Berkheyopsis angolensis. on the geochemistry of the catchment area and certain geomorphological features. T h e vegetation varies greatly 3. Riparian bushland fringing the Bero and Flamingos in relation to the amount of water available and its rivers is characterized b y Tamarix usneoides, Cordia salinity. sinensis and Euclea pseudebenus. Associates include: In small erosion channels with non-brackish water one finds Citrullus ecirrhosus and a few annual herbs Sporobolus robustus, A triplex halimus, Lotus arabicus (mossamedensis), Arthrocnemum indicum, Psoralea obtu(Cleome, Tribulus). sifolia and Asthenatherum {Danthonia) mossamedense. W h e n the amount of storm-water increases certain shrubs such as Adenolobus pechuelii, Parkinsonia afri4. O n well-drained gravel soils near the city of cana and Commiphora saxícola {dulcís) appear. Mossamedes there are communities of Euphorbia virosa O n the margins of larger rivers taller shrubs and {dinteri) associated with species of Aristida, Stipagrostis, small trees, such as Rhus lancea, Salvadora pérsica, Ficus and Eragrostis. sycomorus, Euclea pseudebenus, Acacia erioloba and A. albida occur in riparian woodland. 5. Sarcocaulon mossamedense is dominant in the dry In the bed of the lower Kuiseb on sand dunes with rocky zone at the north of the Reserve. Other species are subterranean water, Acanthosicyos horridus, a leafless virtually absent owing to the strong prevailing winds. gourd with green thorns, forms dense impenetrable Salvadora pérsica occurs in saline depressions. thickets. W h e r e the water is brackish the salt-secreting 6. B y far the most widespread type is a grassland Tamarix usneoides occurs with Lycium tetrandrum. dominated by desert annuals with scattered individuals In river estuaries, flood plains with accessible ground of Welwitschia bainesii. The largest plants of Welwitschia water support the following halophytes: Zygophyllum have crowns 1 m or m o r e in diameter and leaves up to stapfii, Arthraerua leubnitziae, Salsola spp., Suaeda 2 m long. They are found near the coast, where they are plumosa and A rthrocnemum dunense. the only conspicuous feature of the vegetation, and are Springs with slightly brackish water are surrounded scattered about the arid plain at intervals of 50 to by communities of Phragmites, Odyssea (Diplachne) pau100 metres. Lichens are locally plentiful. Except after cinervis and Cyperus laevigatus. falls of rain, other species are not apparent, except for the dead remains of annual species of Stipagrostis and other plants. Further inland the Welwitschia plants are The desert of Mossamedes smaller and the associated vegetation becomes more Refs.: Barbosa (1970, p. 251-61); de Matos & de Sousa (1970); luxuriant. Within the desert area, the latter consists Diniz (1973, p. 269-90); Whellan (1965). largely of grass which forms quite a sward and is Phots.: Barbosa (1970: 28.3, 29.1-3); de Matos & de Sousa" probably the main food of the Springbok and other (1970: 1-3, 6). antelopes, which are fairly numerous (Whellan, 1965). Bushes and dwarf trees are rare and local, and occur The N a m i b desert continues into the south-western as scattered individuals in water-receiving depressions in corner of Angola to a short distance north of the more humid eastern parts. In the western part of the Mossamedes. Rainfall is less than 100 m m per year, but Reserve, Welwitschia occurs o n gravel soils and is most atmospheric humidity is high. T h e flora is a conabundant along drainage lines. Its principal associates tinuation of that of the N a m i b proper, and the are: Stipagrostis subacaulis, S. hirtigluma, Eragrostis vegetation types are similar, but species of tropical porosa, Enneapogon cenchroides, Tricholaena monachne, affinity are m u c h more numerous than further south.
The Karoo-Namib regional centre ofendemism
Dicoma foliosa, Indigofera teixeirae, Geigeria spinosa, Hibiscus micranthus, Aloe littoralis, Sarcocaulon mossamedense, Sesuvium portulacastrum, Lophiocarpus polystachyus and Lotononis tenuis. Further to the east the following grasses join the assemblage listed above: Aristida hordeacea, Stipagrostis hochstetterana, Danthoniopsis dinteri, Tetrapogon tenellus and Rhynchelytrum repens (villosum). Further east still, towards the edge of the desert, Welwitschia occurs in depressions with the microphanerophytes, Acacia tortilis, A. reficiens subsp. reficiens, and Maerua angolensis. Other associates include Lycium decumbens, Hoodia currorii, Monsonia senegalensis, Aristida hordeacea, A. rhiniochloa, Stipagrostis uniplumis, S. hochstetterana, Schmidtia pappophoroides and S. kalahariensis. Rocky outcrops in this part of the Reserve support Euphorbia subsalsa, Commiphora sp., Sterculia setigera and Sansevieria cylindrica. Less information is available for other parts of the
145
Mossamedes desert. North of Porto Alexandre, Welwitschia almost reaches the coast, but to the south and extending far into Namibia, a belt of mobile sand dunes 30-50 k m wide creates conditions approaching absolute desert (P. B a m p s , pers. c o m m . ) . Thus, between Espinheira and F o z do Cunene, Welwitschia drops out at k m 50 (its southern limit in Angola), and, apart from a few cactiform euphorbias and rare individuals of Zygophyllum orbiculatum sheltered by rocks, there are no plants until the halophytic communities of the coast are reached (P. B a m p s , pers. c o m m . ) . Further north between Porto Alexandre and lona, Welwitschia does not appear until k m 49 where it occurs in sand dunes with Acanthosicyos. At the m o u t h of the Cunene, halophytic communities include Cotula coronopifolia, Heliotropium curassavicum, Samolus valerandi, Chenopodium ambrosioides, Tetragonia reduplicata and Cyperus laevigatas. T h e sand dunes near by are devoid of perennial plants except for a few individuals of Rhigozum angolense and Phyla cf. nodiflora (P. B a m p s , pers. c o m m . ) .
vu
The Mediterranean regional centre of endemism
Area, geographical position, geology and physiography Climate Flora Mapping units Vegetation Mediterranean forest Mediterranean broad-leaved sclerophyllous forest Quercus ilex sclerophyllous forest Querela suber sclerophyllous forest Quercus coccifera sclerophyllous forest Mediterranean coniferous forest Juniperus phoenicea forest Cupressus sempervirens and C. atlántica forest Tetraclinis articulata forest Pinus halepensis forest Pinus pinaster forest Cedrus atlántica forest Abies pinsapo and A numidica forest Juniperus thurifera forest Mediterranean deciduous forest Quercus fa*ginea forest Quercus pyrenaica forest Quercus afares forest Mediterranean bushland and thicket Mediterranean shrubland Altimontane Mediterranean shrubland Secondary Mediterranean shrubland (maquis and garrigue) Mediterranean anthropic landscapes
Area, geographical position, geology and physiography This section deals only with the African part of the Mediterranean Region, the Maghreb. Essentially it is the region of folded mountains at the north-western extremity of the continent. (Area: 330000 k m 2 . ) T h e landscape is dominated b y the Atlas mountains, which are largely the product of Tertiary folding and uplift of sediments deposited over a long period in the ocean which lay between the African and Tyrrhenian shields. T h e various ranges of the Atlas are separated b y plateaux and basins. Coastal lowlands occupy a relatively small area. T h e Atlas mountains extend for over 3000 k m from north M o r o c c o to Tunisia. T h e y trend from W S W . to E N E . and run roughly parallel to the Mediterranean coast. T h e y are best developed in Morocco, where M o u n t Toubkal reaches an altitude of 4165 m in the High (Great) Atlas, a mountain range with m a n y snowcapped peaks. In Algeria altitudes d o not exceed 2500 m and in Tunisia 1500 m . T h e oldest of the fold mountains, the Rif Atlas, forms a coastal range which extends from south-east of Tangier to the M o l o u y a river valley and continues in Algeria as the Tell Atlas. In s o m e places there is a separate coastal range, the Maritime Atlas, between the Tell Atlas and the sea. T h e High Atlas extends from the Atlantic coast near Agadir, and in Algeria becomes the Sanaran Atlas, which rarely exceeds 2 0 0 0 m and mostly lies in the Sahara/Mediterranean transition zone. In eastern Algeria the Saharan Atlas a n d the Tell Atlas approach each other. Here occurs Algeria's highest peak, the 2328 m high Djebel Chélia in the Aurès M t s , which are structurally part of the Saharan Atlas. In M o r o c c o the Middle Atlas, which diverges from the High Atlas in a north-easterly direction, consists mainly of a plateau bordered b y mountain chains o n the south and east. T h e Anti-Atlas, which for most of its length is in the Mediterranean/Sahara transition zone, is an elevated part of the African shield. It has a tabular surface at about 1500 metres, but its highest peak is nearly 3900 m above sea-level. It is joined to the High Atlas b y the volcanic formations of Djebel Siroua (3304 m ) , but
The Mediterranean
regional centre of
further west the two ranges are separated by the Souss alluvial plain, which occupies a structural depression. The Atlas ranges extend into Tunisia as the Northern Tell, the High Tell and the L o w Tell. The lithology of the Mediterranean Region is diverse. The prevalent rocks are sediments, sometimes metamorphosed, of Triassic, Jurassic, and Cretaceous age, especially limestones. More recent Upper Tertiary and Quaternary deposits are relatively restricted. There are also small exposures of the Precambrian basem*nt, and outcrops of volcanic rocks.
Climate Most rain falls in winter, and nearly everywhere is between 250 and 1000 m m per year. The summer is hot and dry and is more extreme than that of the Cape Region. Frost is widespread but some parts of the coastal lowlands are frost-free, whereas parts of the interior m a y experience frost for up to seven months each year. In the high mountains snow frequently lies for long periods. (See Fig. 13.) The relationships between climate and vegetation in the Mediterranean Region have been studied by several authors w h o have defined various climatic indices. Those of Emberger (Emberger, 1955a; Sauvage, 1961, 1963) and Bagnouls and Gaussen (1957), which are the best known, are summarized by Quézel (1976). The method of Bagnouls and Gaussen provided the basis of a bioclimatic m a p of the Mediterranean zone (UnescoF A O , 1963). The classification of Emberger, however, has been more widely used in North Africa and his1 étages bioclimatiques are frequently referred to in the following pages. Emberger defines four main 'étages' using a climatic index based on mean annual rainfall, mean m i n i m u m temperature of the coldest month, and mean m a x i m u m temperature of the warmest month. They roughly correspond, however, to annual precipitation as follows: — arid 'étage': 300-500 m m per year; — semi-arid 'étage': 500-700 m m per year; — subhumid 'étage': 700-1000 m m per year; — humid 'étage' > 1000 m m per year. For each 'étage' there are five variants based on the m e a n m i n i m u m temperature of the coldest month, as follows:
— hot>7°C; — — — —
temperate: 3°-7°C; fresh: 0°-3°C; cold: minus 5°-0°C; very cold: < minus 5° C .
1. In French ecological literature the word étage is used in two senses, either to designate strictly altitudinal vegetation belts, or to designate areas which can be characterized in terms of climate and vegetation, but of which the spatial relationships are more often of the checkerboard type than strictly altitudinal. It is the latter sense which has been adopted in North Africa by Emberger and others.
endemism
147
Emberger also recognized a Mediterrano-Saharan 'étage' and a Mediterranean high-mountain 'étage'. The latter largely corresponds to the very cold variants of the 'étages' Usted above. The Sanaran 'étage' is represented in the Mediterranean Region sensu stricto only by very small enclaves.
Flora About 4000 species occur in the north African part of the Mediterranean Region (excluding the Mediterranean-Sahara transition zone). O f these c. 72.5 per cent are Mediterranean endemics, though only c. 20 per cent of them are confined to North Africa. Endemic families. None. Aphyllanthes (monotypic, Liliaceae) is sometimes given family rank. Globulariaceae, which has most species in the Mediterranean Region, is also represented by two species of Poskea in Somalia and Socotra. The Cneoraceae is confined to the Mediterranean Region, the Canaries and Cuba. Endemic genera. About 250 genera have their greatest concentration of species in the Mediterranean Region, but m a n y are not strictly endemic. Thus, among genera occurring in North Africa, Cyclamen reaches Persia, and Cistus extends from the Canary Islands to TransCaucasia. Other near-endemic genera include Ceratonia, Helianthemum, Genista, Lavandula and Ononis. More strictly endemic are Anagyris, Chamaerops, Coriandrum, Halimium, Spartium, Tetraclinis and m a n y genera of Cruciferae. Endemic species. Non-endemic genera with more than 50 endemic species in Mediterranean North Africa include: Astragalus, Centaurea, Euphorbia, Linaria, Silène, Teucrium, Trifolium and Vicia. M a n y of the most characteristic Mediterranean species, including Arbutus unedo, Cedrus atlántica, Laurus nobilis, Myrtus communis, Nerium oleander, Quercus coccifera, Q. ilex, Q. suber and Vitex agnus-castus, are endemic, but belong to non-endemic genera. Linking elements. Approximately 1.5 per cent of the species occurring in Mediterranean North Africa sensu stricto are cosmopolites; 20 per cent, including Acer campestre, Alnus glutinosa, Betulapéndula (fi. alba auct.), Calluna vulgaris, Carex capillaris, Digitalis purpurea and Prunus padus, are Boreal linking species. Only 3 per cent, including Lupinus varius (pilosus). Retama retam, Stipagrostis (Aristida) pungens and Ziziphus lotus are Sanaran linking species, and 2.2 per cent are IranoTuranian linking species. Other linking species include Erica arbórea, which also occurs on the high mountains of the Sahara and East Africa, and Pistacia atlántica, which extends from the Canary Islands to Afghanistan.
Vegetation of the floristic regions
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The Mediterranean regional centre of endemism
Floristic relationships with the Cape Region are slight (Burtt, 1971). Only 30-35 Mediterranean species, of which 15 are in Erica, belong to characteristic Cape genera.
Mapping units 10. Mediterranean sclerophyllous forest. 23. Mediterranean montane forest and altimontane shrubland. 78. Mediterranean anthropic landscapes.
Vegetation Most of the Maghreb was formerly covered with forest, but o n clay soils in the semi-arid 'étage' scrub forest dominated by Olea europaea and various types of bushland or thicket m a y formerly have represented the climax. Otherwise, non-forest woody vegetation w a s confined to shallow soils, wind-swept ridges and coastal habitats, and the summits of the higher mountains. Ionesco and Sauvage (1962) recommend the use of the Spanish word 'matorral' to cover all w o o d y types of non-forest vegetation in the Mediterranean Region. This usage, which corresponds to the English 'scrub' (bushland plus shrubland) has m u c h to recommend it. Although grasses are well represented in the Mediterranean flora, climax grasslands were formerly of limited extent. Grasslands dominated by Stipa tenacissima and Lygeum spartum, which are probably largely secondary, are found in the drier parts of the Mediterranean Region. They are more characteristic, however, of the Mediterranean/Sahara transition zone and are described in Chapter XVIII. Ampelodesma mauretanicum also locally forms a secondary 'steppe' grassland in higher-rainfall areas where heavy cutting and grazing o n steep clay slopes have led to soil removal. Communities dominated by Argania spinosa, Acacia gummifera and succulent Euphorbias, which represent the most characteristic vegetation types of the western end of the Mediterranean/Sahara transition zone, also occur very locally in the Mediterranean Region in Morocco, but only as small enclaves or marginal intrusions. They too are described in Chapter XVIII.
149
articulata, dominate coniferous forest, and three species, Quercus fa*ginea, Q. pyrenaica, and Q. afores, dominate deciduous oak forest. O f the remainder, the majority are Mediterranean endemics, or, if they occur elsewhere, their distributions are centred o n the Mediterranean basin. They include: Acer monspessulanum, Arbutus unedo, Celtis australis, Ceratonia siliqua, Chamaerops humilis, Crataegus azarolus, Fraxinus angustifolia, F. xanthoxyloides, Juniperus oxycedrus, Laurus nobilis, Olea europaea, Phillyrea angustifolia (including P. latifolia and P. media), Pinus pinaster, Pistacia atlántica, P. lentiscus, P. terebinthus, Prunus lusitanica, Pyrus gharbiana, P. cossonii (longipes), P. mamorensis and Rhus pentaphylla. Several Eurosiberian linking species also occur in Mediterranean North Africa, notably Acer campestre, Alnus glutinosa, Betula péndula, Crataegus monogyna, Ilex aquifolium, Populus trémula, Prunus avium, P. padus, Sorbus aria, S. domestica, S. torminalis, Taxus baccata and Ulmus campestris. All have restricted distributions and are virtually confined to the humid and subhumid 'étages' of the Rif in Morocco, the wetter parts of the coastal plain and coastal ranges in Algeria, and the wetter parts of the seaward slopes of the Atlas M t s . Most types of Mediterranean forest in North Africa have suffered extreme degradation nearly everywhere, and have disappeared completely from large areas. S o m e are represented today b y n o more than tiny relictual stands. It is possible that some types have gone completely except for a few relict individual trees. Three such types are briefly mentioned below. 1. Celtis australis is one of the tallest deciduous trees in Mediterranean Africa. It reaches a height of 25 m . Today it occurs as rare individuals in the lowlands and is even more scattered inland, where it ascends to 1300 m . According to Monjauze (1958) this species has suffered more at the hands of m a n and hisflocksthan any other Mediterranean tree. Monjauze presents evidence which suggests that, o n deep soils in the subhumid 'étage' and in the lower wanner part of the humid 'étage', Celtis australis formerly dominated forests which had an understorey of Laurus nobilis and a herb layer which included Acanthus mollis.
2. Pistacia atlántica is also a fine deciduous tree which reaches a height of 20 m and a diameter of 1 m . It has a Mediterranean forest scattered distribution from the Canary Islands to (mapping units 10 and 23) Afghanistan. In the Maghreb it is one of the most widely Most of the North African part of the Mediterranean distributed species but occurs as isolated individuals, not Region sensu stricto w a s formerly covered with forest. in forest stands. It is most plentiful in the w a r m semiOnly small relicts survive, but they include at least arid 'étage' and ascends to 2000 m in the west and 60 tree species, of which 16 are major dominants. Three 3000 m in the east. It regenerates freely from seed, species, Quercus ilex, Q. suber and Q. coccifera, dominate especially in the protection of clumps of Ziziphus lotus. evergreen sclerophyllous forest. T e n species, Abies Its rarity today is due to its susceptibility w h e n young to numidica, A. pinsapo subsp. marocana, Cedrus atlántica, browsing and fire, and to the limited occurrence of deep Cupressus atlántica, C. sempervirens, Juniperus phoenicea, soil because of extensive erosion. Monjauze (1968) J. thurifera, Pinus halepensis, P. pinaster and Tetraclinispostulates that on deep soils in the semi-arid 'étage' the
150
Vegetation of the floristic regions
climax forest was formerly dominated b y Pistacia atlántica mixed with sclerophyllous oaks.
Quercus ilex is essentially a species of the mountains. In the Rif it ascends to 2200 m , and is very widespread at higher altitudes above the Tetraclinis, Quercus suber and Pinus halepensis forests of the lower slopes. In the 3. In marshy places on the Algerian coast the climax Middle and High Atlas it covers an enormous area forest was probably composed of Ulmus campestris between 600 and 2900 m on the Atlantic slopes, but is mixed with Fraxinus angustifolia, Populus alba, Salix m u c h less abundant o n the Sanaran slopes and o n alba and Laurus nobilis. Mediterranean slopes of pronounced relief. In the AntiAtlas it forms forest only at the western end. The forests of Morocco are floristicallyricherand more T h e ecology of Quercus ilex is diverse. It occurs diversified than those occurring elsewhere in Mediprincipally under temperate and cold climates in the terranean Africa, and the following account is based on semi-arid, subhumid, and humid 'étages'. In Morocco, them, but the forests of Algeria and Tunisia are despite the fact that the forests have been decimated essentially^ similar and the dominants are mostly the over large areas, Quercus ilex has survived in anthropic same. Quercus pyrenaica and Abies pinsapo, subsp. landscapes more frequently than other Mediterranean marocana, however, are confined to Morocco, and are forest trees because of its resistance to fire and ability to replaced by Q. afares and A . numidica in Algeria; neither sucker from stumps and damaged roots. It occurs on a occurs in Tunisia. Cedrus atlántica and Pinus pinaster are wide range of soils but is absent from most clayey also absent from Tunisia, and Cupressus sempervirens, depressions. sensu lato, is absent from Mediterranean Algeria. Five forest dominants, Quercus ilex, Q. coccifera, Towards its lower altitudinal limits in the hot semiPinus halepensis, Cupressus sempervirens and Juniperus arid 'étage' it is in contact with Tetraclinis articulata, phoenicea, also occur in Cyrenaica. Their former Juniperus phoenicea and Pinus halepensis and, very distribution w a s restricted and only small degraded locally, with Quercus suber. In Morocco it occurs in the remnants remain today. Tetraclinis articúlala also occurs lowlands of the semi-arid 'étage' only south of Rabat in Cyrenaica but is doubtfully native. where, on siliceous soils, it forms a band between the forests of Quercus suber and those of Tetraclinis. These Argania spinosa is extremely localized in the forests are low and open, and are formed of stunted Mediterranean Region sensu stricto but forms extensive twisted trees. They differ from the nearby Quercus suber communities at the western end of the sub-Mediforests only in their dominant species. T h e shrub and terranean transition zone. They are described in field layers of the two forest types are virtually identical. Chapter XVIII. This is because during the wet season their climates are Because m a n y of the dominants of Mediterranean similar. During the dry season the Quercus ilex forests forest grow within a wide range of climate, classification experience hotter and drier conditions. It is summer is difficult. In the following account the distribution of drought which eliminates Q. suber. the main types is described in outline in relation to the 'étages' bioclimatiques of Emberger (page 147). Towards the upper limit of its range in the cold semiarid 'étage', Quercus ilex is in contact with Juniperus phoenicea, Juniperus thurifera and Cupressus atlántica. Mediterranean broad-leaved sclerophyllous forest In the subhumid and humid 'étages', where it attains its m a x i m u m development, it occurs in mosaic with, and This type occupies about half of the total forest area. sometimes intermingled with, Quercus suber, Q. fa*ginea, Everywhere it is dominated by Quercus ilex, Q. suber or Q. pyrenaica, Pinus pinaster and Cedrus atlántica. Q. coccifera, the distributions of which are almost Under favourable conditions Q. ilex attains a girth mutually exclusive. B y far the largest area (2100000 ha, of 3 m and a height of 20 m with a magnificent wideor more than two-thirds of the total) is Q. ilex forest. spreading crown. M o r e frequently, however, it is a Q. coccifera is relatively restricted (44000 ha). smaller tree with a short trunk and more compact crown. Its coppice sometimes forms thickets, in which seedQuercus ilex sclerophyllous forest lings of Cedrus, Pinus halepensis and P. pinaster find Refs.: Boudy (1948, p. 139-40; 1950, p. 299-351); Emberger protection from browsing animals and can regenerate. (1939, p. 107-10, 111-14, 135-6); Métro (1958, p. 68-73); T h e ability of Quercus ilex to withstand competition Peyerimhoff (1941, p. 53); Quézel(1976). is remarkable. It is capable of persisting as suppressed Phots.: Boudy (1948: 6-8; 1950: 38-46); Emberger (1939: 9.2); individuals beneath the thick canopy of Cedrus forest. Métro (1958: 4). W h e n a cedar dies, Quercus ilex grows rapidly to occupy the vacant space, but regenerating cedars ultimately Quercus ilex is the most widely distributed and most overtop it and cast their shade again. abundant tree in Mediterranean Africa, and probably in the Mediterranean basin generally. In the Maghreb it is virtually absent from the lowlands below 400 m , but In the subhumid 'étage' forms forests above that altitude up to 2400 m in the Nearly all the surviving stands of Quercus ilex forest in Atlas. Scattered individuals ascend to the tree line at c. 2900 m . Morocco occur in this 'étage'. They cover enormous
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areas o n the lower slopes of the Rif and the Atlantic slopes of the Middle Atlas and the High Atlas. In the Rif Q. ilex plays an important part only on the Mediterranean slopes, where it occurs both on calcareous and siliceous soils. O n the moister Atlantic slopes it is replaced by other species of Quercus, especially Q. suber, except on calcareous soils, which are of limited extent. All the stands of subhumid Q. ilex forest in the Middle Atlas are on calcareous soils. Those in the High Atlas occur both on calcareous and siliceous soils. At the western extremity of the High Atlas and in the mountains north of Essaouira (Mogador), Q. ilex forms islands of forest above 650 m , at which altitude it replaces Tetraclinis. T h e southernmost stand of subhumid Quercus ilex forest is in the massif of Kest in the western Anti-Atlas. A little further to the south-west, scattered individuals of Q. ilex occur in populations of Tetraclinis on the highest peaks of Ifni (1250 m ) . Subhumid Quercus ilex forest has a low, dense, almost closed canopy w h e n it is intact; the underwood is sparse. Such stands are rare. Most have been degraded and invaded by heliophilous species. Emberger (1939) recognizes four types of subhumid Quercus ilex forest based on temperature and substrate, but their floristic characterization is somewhat weak.
Together with the Atlas Cedar (Cedrus atlántica), the Cork O a k is the most valuable tree in North Africa. Unlike Q. ilex it is confined to the western half of the Mediterranean basin and does not extend further east than Italy. In the Maghreb it occupies 843000 ha. T h e canopy of Q. suber forest is usually low and somewhat open, and occurs at a height of 6-12 m in the drier types, but is closed and taller (15 m ) in the wetter types. There is usually a well-developed shrub layer, 2-4 m tall, except w h e n the canopy is dense and continuous; then the underwood is less abundant or disappears. Q. suber avoids wet, cold conditions. In Morocco it occurs locally almost at sea-level and ascends to 1600 m in the Rif, 1550 m in the Middle Atlas, and 2100 m in the High Atlas. A t different altitudes its climatic and edaphic requirements are somewhat different. Like Q. ilex it occurs in the semi-arid, subhumid and humid 'étages'. A t the lowest altitudes o n clay soils it is replaced by Olea-Pistacia lentiscus scrub, on calcareous soils by Pinus halepensis, and o n the littoral by P. pinaster or Juniperus phoenicea. In the mountains it is in constant competition with Q. fa*ginea, and in Algeria with Q. afares, which are better adapted to cold, grow taller, form a more continuous canopy, and regenerate more prolifically. Where Q. suber and the deciduous oaks occur in mixed stands the latter would be dominant if it were not for the effects of fire, to which they are less In the humid 'étage' well adapted than Q. suber. At higher altitudes and o n This type of Q. ilex forest differs from drier types in its drier soils or where the dry season is severe, Q. suber is greater stature and denser canopy, the abundance of replaced by Q. ilex. Q. suber never forms forests o n bryophytes and lichens, and in the deep, humus-rich calcareous soils. soil. FloristicaUy it shares several species with the forests Semi-arid Q. suber forest covers extensive areas in of the Eurosiberian Region. In Morocco it is confined to N W . Morocco but in Algeria it occurs only as relics in the Rif and the Middle Atlas. the Oran-Mascara region. Nearly all the Q. suber forests T h e best-known examples occur near Azrou in the in Algeria and Tunisia are of the humid and subhumid Middle Atlas. T h e canopy is almost pure but there are types. In Morocco, these wetter types occur only in the scattered individuals of Acer monspessulanum, T'axus Rif. There are certainly some floristic differences baccata and Sorbus torminalis. In the underwood m a n y between the semi-arid, subhumid, and humid types but species occur including: Coronilla valentina {glauca), superimposed o n this are floristic differences which d o Cotoneaster fontanesii, Crataegus monogyna, Cytisus not appear to reflect ecological conditions. battandieri, Daphne laureola, D . gnidium, Ilex aquifolium, Except in the remotest places, the bark of Q. suber is Juniperus oxycedrus, Lonicera etrusca, Rosa sp., Rubus stripped from the boles of all mature trees every nine ulmifolius, Ruscus aculeatus, Viburnum lantana and V. years to provide the cork of commerce. tinus. Climbers are represented by Asparagus acutifolius, Clematis cirrhosa, Hederá helix, Rubus, Smilax áspera, In the semi-arid 'étage' and Tamus communis. In ravines one encounters Euonymus latifolius, Ligustrum vulgare, Rhamnus This type is a xerophilous open forest with a floristically catharticus, Salix cinérea and Sorbus aria. T h e diversified poor underwood but with a field layer rich in therofield layer includes m a n y geophytes but annuals by phytes. It occurs principally o n Pliocene sands or o n contrast are very rare w h e n the forest is not degraded. schist.
Quercus suber sclerophyllous forest Refs.: Boudy (1948, p. 137-9; 1950, p. 29-180); Emberger (1939, p. 101-5, 118-19, 136-7); Métro (1958, p. 51-68); Peyerimhoff (1941, p. 54-5); Quézel (1976); Sauvage (1961). Phots.: Boudy (1948: 4-5; 1950: 2-25); Métro (1958: 3).
T h e best-known example in Morocco is the M â m o r a forest which occurs o n deep sands. A n endemic Pear, Pyrus mamorensis, is scattered in the canopy. In most of the forest the shrub layer is dominated b y Cytisus (Teline) linifolius, and in the eastern part by Halimium halimiifolium. In the more open parts Cytisus arboreus, Daphne gnidium, Lavandula stoechas and Ulex (Stawacanthus) boivinii are found. Halimium libanotis is
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abundant in the western part of the forest. Cistus salviifolius is c o m m o n only in those parts which are moist in winter. Chamaerops humilis is especially abundant in places where there is a clay horizon near the surface. In large clearings and at the edge of the forest Thymelaea lythroides and the conspicuous composite Ormenis multicaulis are particularly c o m m o n . A t the centre of large clearings the w o o d y plants have often disappeared and are replaced by open seasonal herbaceous vegetation, in which bulbous plants, especially Urginea marítima, Dipcadi serotinum, Asphodelus microcarpus and A . aestivus, are abundant. In this extremely degraded community regeneration of Q. súber is sporadic or absent.
Quercus coccifera sclerophyllous forest Refs.: Boudy (1950, p. 378-81); Emberger (1939, p. 115); Métro (1958, p. 101): Peyerimhoff (1941, p. 53-4).
T h e Kermes O a k , Quercus coccifera (including Q. calliprinos), has a patchy circum-Mediterranean distribution. In the Maghreb it occupies 44000 ha. In Morocco, it occurs only in the Rif, except for a single station further south. Towards the east, in Algeria and Tunisia, it is m u c h more abundant and occurs on coastal sands and extends a short distance inland, where it is replaced by Q. suber. Q. coccifera grows on a very wide range of soils, both acid and alkaline, and under a m e a n annual rainfall from 450 to m o r e than 1000 m m . Q. coccifera is usually seen as a dense, m u c h branched shrub, but locally it is a tree, as in the mountains above Ceuta in Morocco, where it occurs In the subhumid 'étage' with Taxus baccata. Here it is possible for a m a n to walk unimpeded beneath its boughs. It is possible that Q. T h e shrub layer is better developed than in the humid coccifera formerly formed extensive forests. Almost type. Below 1200 m in the Rif Q. suber forms a canopy everywhere it has been cut for charcoal, but, because of at 15 m beneath which there is a dense shrub layer 4-5 m its capacity to sucker and withstand repeated cutting, it tall of Arbutus unedo and Erica arbórea. Beneath this still covers large areas in its familiar shrubby form. In a double canopy there are scattered small shrubs such as Cistus salviifolius, Cytisus monspessulanus and Lavandula few places in Algeria it can still be seen in relict patches of relatively undisturbed forest as a tree up to 12 m tall stoechas. In the meagre field layer Carex distachya and with a bole about half that length (P. J. Stewart, pers. Eryngium tricuspidatum are found. If a tree dies it is c o m m . ) . Here, on maritime sands, it is often associated eventually replaced but larger gaps in the canopy are with 8-9 m tall Juniperus phoenicea. A t Mostaganem in filled b y a very dense growth of Arbutus and Erica Algeria, Pistacia lentiscus, Olea europaea, Phillyrea arbórea, which seriously hinders the regeneration of the angustifolia, Ephedra altissima and Ceratonia siliqua (on Cork O a k . Repeated cutting or burning results in the the landward side only) occur as shrubs or occasionally invasion and eventual dominance of heliophilous shrubs such as Cistus populifolius, C. crispus, Halimium lasioca- as small trees in the understorey. Small shrubs are: Retama monosperma, Calicotome villosa {intermedia), lycinum, Erica umbellata and Calluna vulgaris. Further Withania frutescens, Clematis cirrhosa, Asparagus acutidegradation, as in the humid zone, is followed by the folius, Lycium intricatum, Lavandula dentata and establishment of even more xerophilous species, which Teucrium polium (P. J. Stewart, pers. c o m m . ) . are otherwise confined to the semi-arid 'étage'. In the humid 'étage'
Mediterranean coniferous forest
Nearly half of the area occupied b y Mediterranean In Morocco only a few undisturbed stands survive in the forest in the Maghreb is dominated by coniferous forest. Bab-Azhar forest in the Rif between 1200 and 1500 m , There are 10 main dominants: Abies numidica, but this type occurs extensively in eastern Algeria and A . pinsapo subsp. marocana, Cedrus atlántica, Cupressus Tunisia. Beneath the closed canopy the shrub layer is atlántica, C. sempervirens, Juniperus phoenicea, J. sparse and consists of scattered individuals of Cytisus thurifera, Pinus halepensis, P. pinaster and Tetraclinis villosus (triflorus), C. maurus, Cistus salviifolius and Ulex articulata. The areas they occupy vary greatly from a few boivinii. The field layer, which is also exiguous, includes hundred hectares (A. numidicá) to 1300000 ha (Pinus Dactylis glomerata and Pteridium aquilinum. If a small halepensis). gap in the canopy is formed by the death or removal of a tree the above-mentioned shrubs increase in abundance Four other conifers occur in the Maghreb, but not as and are joined b y more heliophilous species. Y o u n g dominants. Thus: plants of Q. suber can establish themselves in such gaps — Taxus baccata is frequently associated with Abies but only outside the area occupied b y the roots of numidica, A . pinsapo, Cedrus atlántica or Quercus ilex in established trees, which would inhibit their growth. the humid 'étage'. — Pinus nigra is only k n o w n from two localities, one H u m i d Q. suber forest differs from the subhumid in Algeria, where it occurs in the middle of Cedrus type in the presence of Alnus glutinosa, Prunus avium, forest, and the other in the Rif in Morocco where it is Quercus fa*ginea and Taxus baccata in valley bottoms. mixed with Pinus pinaster. — Juniperus oxycedrus occurs throughout the Maghreb, either as a tree 10 m tall with a bole 1 m in
The Mediterranean regional centre ofendemism
diameter, or more often as a smaller bushy plant. It ascends from sea-level to 3000 m and is nearly always associated with Quercus ilex or Juniperus phoenicea. — T h e Holarctic species, Juniperus communis, forms dense cushions above the timber-line.
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Tetraclinis and Quercus ilex it has n o floristic individuality. In the lower part its associates are the most tolerant m e m b e r s of the former and in the upper part of the latter. / . phoenicea only occurs at two places in theRif.
Juniperus phoenicea forest Refs.: Boudy (1948, p. 134-5; 1950, p. 741-53); Emberger (1939, p. 78-86); Métro (1958, p. 77-8); Peyerimhoff (1941, p. 50). Phot.: Boudy (1950: 116). Juniperus phoenicea extends from the Canary Islands to Arabia and Jordan. It has two principal habitats, coastal sands and the high plateaux and mountains of the interior. It is almost confined to the semi-arid 'étage' in which it occurs on a variety of soils. In Morocco it often occurs in a zone between Tetraclinis forest and Quercus ilex forest, but in the coldest parts of the semi-arid 'étage' it replaces Tetraclinis. In Algeria it is often mixed with Pinus halepensis, but it is in the Sanaran Atlas bordering the desert that it achieves its greatest extension. Today, Juniperus phoenicea usually occurs as a bushy tree less than 7 m tall in open stands, which physiognomically are wooded grassland. They probably represent degraded forest. This is suggested by the fact that it can occur as a tree 8-9 m tall (P. J. Stewart, pers. c o m m . ) with a massive trunk u p to 2 m in girth. Although J. phoenicea locally dominates thicket o n coastal sands where it is exposed to strong wind, or bushland in the interior where the soils are too shallow to support forest, it is likely that formerly it also dominated short forest which covered considerable areas. Emberger, for instance, states that the dunes near Essaouira (Mogador) were once covered with J. phoenicea forest. The fact that 'tout M o g a d o r est construit avec du bois de Juniperus phoenicea' testifies to its former abundance as a relatively large tree. For Morocco, Emberger describes the following three types of/, phoenicea communities.
3. Trans-Atlas communities These are the driest types and are very degraded, often almost to the point of total disappearance. They occur on the southern slopes of the Great Atlas and AntiAtlas and on the lower slopes of the Upper Moulouya Valley. They occupy a zone above Stipa tenacissima grassland which m a y be, at least in part, secondary (page 229). O n this side of the Atlas, Q. ilex forests formerly occupied only a narrow band, and, at least locally, with increasing altitude, / . phoenicea is in contact with J. thurifera. The more prominent associates of Juniperus phoenicea in these degraded communities include: Fraxinus xanthoxyhides, Buxus baleárica, Rhamnus alaternus, R. oleoides, Adenocarpus bacquei, Carthamnus fruticosus, Genista myriantha, Globularia alypum, Lavandula multifida, Artemisia herba-alba and Stipa tenacissima.
Cupressus sempervirens and C . atlántica forest Refs.: Boudy (1950, p. 764-70); Destremau (1974, p. 67-76); Emberger (1939, p. 100); Métro (1958, p. 79); Peyerimhoff (1941, p. 49-50). Phots.: Boudy (1950: 122-3).
Cupressus sempervirens s.l. is a circum-Mediterranean species which extends as far east as Jordan. In the Maghreb it is indigenous only in two principal localities, namely the Mactar region of central Tunisia, and in Morocco, in the basin of O u e d N'fis, south of Marrakech, where it occupies 10000 ha between 1100 and 1800 m . There are also a few small scattered populations in the High Atlas between 1000 and 2000 m . C. sempervirens has been so widely planted since 1. Littoral communities classical times that there is some doubt concerning the extent of its natural distribution. T h e fastigiate form, /. phoenicea only grows well under the protection of the var. sempervirens, which probably does not occur in the first ridge of dunes. At Mehdia on the Atlantic coast the wild, is absent from the Tunisian population, which is tiny remaining fragments are in the form of windprobably indigenous (P. J. Stewart, pers. c o m m . ) . T h e trimmed thicket (White, M S , 1974). They contain Phillyrea angustifolia, Pistacia lentiscus, Ephedra fragilis, Moroccan plant is so different from the naturally occurring C. sempervirens that in the opinion of most Rhamnus alaternus, R. oleoides, Jasminum fruticans, botanists it should be treated as a distinct species, C. Smilax áspera, Clematis cirrhosa, Asparagus albus and atlántica. Osyris sp. Near Essaouira / . phoenicea is associated with The main population of C. atlántica forms an island m a n y of the species which occur at Mehdia, but with in the Juniperus phoenicea horizon between those of southern elements, including Periploca laevigata and Tetraclinis articulata and Quercus ilex in the semi-arid Helianthemum canariense, in addition. 'étage'. T h e associates of Cupressus atlántica and Juniperus phoenicea are virtually identical. 2. Cis-Atlas communities C. atlántica can develop into a fine tree 40 m or more tall but today, a m o n g the older trees, it is mostly mutiThese occur below 2200 m in the Great and Middle lated wrecks that are seen. It has, however, been Atlas. W h e r e J. phoenicea forms a horizon between
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protected for more than thirty years and is reverting to its natural condition. Tetraclinis articulata/oresi Refs.: Boudy (1948, p. 133-4; 1950, p. 706-39); Emberger (1939, p. 71-8); Métro (1958, p. 79-83); Peyerimhoff (1941, p. 49). Phots.: Boudy (1948: 3; 1950: 105-12); Emberger (1939: 5.1); Métro (1958: 7). Syn.: association du Thuja (Callitricetum) (Boudy, 1948); forêt de Thuja de Barbarie (Emberger, 1939); la Callitraie (Emberger, 1939). Tetraclinis articulata is confined to North Africa, except for a small population in Malta and another in the extreme south-eastern corner of Spain. It extends from southern Morocco to Tunisia, with a gap between Algiers and the Tunisian frontier. In Morocco it occurs on the lower Mediterranean slopes of the Rif, and then extends eastwards into Algeria. It also occurs o n the lower northern slopes and in deep rain-shadow valleys of the Middle Atlas and High Atlas and extends round the western end of the latter to the northern slopes of the Anti-Atlas. Extensive Tetraclinis forests occur in the hinterland between Essaouira and Agadir above the Argania scrubland, and in deep valleys of the upper courses ofriversinland from Rabat and Casablanca. Tetraclinis is generally confined to the oceanic and maritime semi-arid 'étages' between sea-level and 1500 m . Cold, especially humid cold, prevents it from ascending higher. It does not occur o n the southern slopes of the High Atlas east of Siroua nor of the Middle Atlas, except at the northern end where the climate is not too continental because of the proximity of the sea. It is found o n both calcareous and siliceous soils, but they are nearly always lithosols. It does not tolerate badly drained conditions. A t the moister limits of its range it is confined to calcareous soils; o n other welldrained soils it is replaced by Quercus suber, and on clay soils by Olea-Pistacia scrubland. A t the moister limits of its range, where it is in contact with Quercus ilex or Q. suber, it usually occupies the xerocline, but at the drier limits, where it intermingles with Argania, it prefers the mesoline. Well-grown Tetraclinis forest is from 12 to 15 m tall, but it is often m u c h shorter than this. T h e narrow crowns form only a light canopy and most of the associated species are heliophilous. Certain species are constantly present in Tetraclinis forest. Others are of m o r e local occurrence. Species which are always present in Tetraclinis forest include Cistus villosus, Ebenus pinnata, Lavandula multifida, Osyris sp. and Teucrium polium, but they are not exclusive. Cistus villosus and Teucrium polium are the least faithful. Other less constant species are Ampelodesma mauritanicum, Anthyllis cytisoides, Brachypodium ramosum, Cistus clusii, Clematis cirrhosa, Ephedra fragilis, Erica multiflora, Genista retamoides, Helianthemum lavanduli-
folium, Jasminum fruticans, Lavandula dentata, Quercus coccifera, Rosmarinus officinalis, Teucrium fruticans (a good indicator of calcareous soil), Viola arborescens (strictly littoral) and the endemic Polygala balansae. Pinus halepensis/oresi Refs.: Boudy (1948, p. 132-3; 1950, p. 639-90); Destremau (1974, p. 5-28); Emberger (1939, p. 94-100); Métro (1958, p. 74-7); Peyerimhoff (1941, p. 48-9). Phots.: Boudy (1950: 90-101); Emberger (1939: 2); Métro (1958: 5). Pinus halepensis, which reaches a height of 20 m , is distributed throughout the greater part of the Mediterranean basin from the southern shores of the Black Sea to Spain and Morocco. It occurs in Cyrenaica. In natural forests it does not extend to the Atlantic seaboard, but grows well w h e n planted there. In southern Morocco the nearest populations to the Atlantic Ocean are 145 k m inland. In North Africa there are immense populations in Tunisia and in South Oran, but they have often been degraded by fire. In Morocco the only extensive forests are in the High Atlas, but there are m a n y scattered smaller stands. T h e total area occupied by Pinus halepensis in North Africa is estimated at 1250000 ha. P. halepensis extends from sea-level to 2000 m , but it is confined to the semi-arid 'etage' and the drier part of the subhumid 'étage'. In the Atlas it does not descend lower than 1200 m . A t the centre of its distribution P. halepensis occurs o n a wide range of soils but towards the edges of its range, where low temperatures or high humidity are limiting, it is confined to calcareous soils. It is often associated with Tetraclinis articulata, Juniperus phoenicea or Quercus ilex. It is less xerophilous than Tetraclinis and does not ascend as high as / . phoenicea. In eastern Morocco it forms islands in Tetraclinis forest, as o n the Melilla peninsula. In this part of its range w h e n P. halepensis forest is degraded it is invaded b y Tetraclinis, which, because of its ability to regenerate vigorously from suckers, is better adapted to recurring fires. In western Morocco the islands of P. halepensis are surrounded b y Q. ilex forest and occur on edaphically favourable sites. Here, where it is at the moister limits of its range, it expands w h e n nearby Q. ilex forest or Cedrus atlántica forest is damaged by fire. P. halepensis occurs on the Sanaran Atlas in Algeria, but not on the Anti-Atlas in Morocco. N o r has it been found on the southern slopes of the High Atlas, but the vegetation there has been terribly degraded by m a n . In Morocco Pinus halepensis forests have n o floristic individuality. P. halepensis normally occurs in mixed forest with Tetraclinis articulata or Quercus ilex, in which the pine is an emergent and its tree associates form the lower canopy. This condition is often brought about by the action of m a n , but some mixtures are probably natural. Thus, on very steep slopes in deep valleys in the High Atlas (White, M S , 1974) P. halepensis occurs as an emergent over an open 'matorral' of Pistacia lentiscus, Quercus ilex, Juniperus phoenicea, J. oxycedrus, Olea
The Mediterranean regional centre of endemism
europaea and Phillyrea angustifolia. These slopes are very unstable. Doubtless some of the contemporary erosion is due to h u m a n influences, but the relief is so accentuated that natural erosion is probably sufficiently active to ensure a permanent habitat for the heliophilous P. hale- i pensis. In Algeria certain species are found more frequently in Pinus halepensis forest than in any other forest type, notably Globularia alypum, Leuzea conifera, Rosmarinus eriocalyx (tournefortii) and, as a forest grass, Stipa tenacissima.
Pinus pinaster forest Refs.: Boudy (1950, p. 691-702); Destremau (1974, p. 29-66); Emberger (1939, p. 115-17,137-8); Métro (1958, p. 48-51); Peyerimhoff (1941, p. 49); Quézel (1976). Phots.: Boudy (1950: 102-4). Pinus pinaster is virtually confined to the western half of the Mediterranean basin. It extends n o further east than Italy and Tunisia. In North Africa its distribution is restricted and it occupies only 28000 ha. It is, however, of considerable, and even more of potential, economic importance. In Morocco P. pinaster grows only in the mountains, in Algeria and Tunisia only o n the coastal plain where it never exceeds an altitude of 700 m . In Morocco it scarcely descends below 1000 m and ascends to 1900 m in the western Rif. O n the northern slopes of the Middle Atlas and the High Atlas it occurs between 1500 and 2200 m . It is possible that the Moroccan and Algerian variants are taxonomically different (Monjauze, Bull. Soc. Hist. Nat. Afr. Nord, vol. 45, 1954, p. 39-54). P. pinaster is confined to the subhumid and humid 'étages'. O n the North African littoral east of Algiers the climate is humid. Rainfall is 1000-1200 m m per year and frosts d o not occur. In the mountains of Morocco it occurs in the humid and semi-humid 'étages'. Rainfall is believed to be 800-1000 m m per year and winter temperature m a y be below zero for long periods. P. pinaster grows on a wide range of soils but is more selective than P. halepensis. In the littoral zone it is found only o n Numidian sandstone. In Morocco it grows both on siliceous rocks and on Jurassic and Cretaceous dolomites, though the soils derived from the latter are often free of calcium carbonate. It always grows o n well-drained soils. A t its upper limit it mingles with Cedrus atlántica and Abies pinsapo. It is rarely associated with P. halepensis. T h e Pinus pinaster forests in Algeria and Tunisia usually contain Quercus suber and Q. fa*ginea. N o n e of the species of the shrub layer, which includes the grass Ampelodesma mauritanicum, is characteristic. All occur in h u m i d Quercus suber forest. In the subhumid Pinus pinaster forests in Morocco, Quercus suber or Q. ilex, according to the nature of the soil, are present in the Rif, and in the Middle Atlas Q. ilex sometimes mixed with Q. fa*ginea usually occurs. In
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the driest forests scattered tufts of Stipa tenacissima occur in the field layer. In the humid P. pinaster forests in the Rif, Abies pinsapo, Cedrus atlántica, Q. ilex or Q. suber are usually present. Cedrus and Quercus ilex are scattered in the canopy of humid P. pinaster forests in the Middle Atlas, which sometimes occur as magnificent, almost pure stands. Cedrus atlántica forest Refs.: Boudy (1948, p. 135-6; 1950, p. 529-611); Destremau (1974, p. 77-90); Emberger (1939, p. 123-31); Métro (1958, p. 34^*6); Peyerimhoff (1941, p. 48); Quézel (1976). Phots.: Boudy (1950: 75-87); Métro (1958: 1). Cedrus atlántica is confined to the mountains of Algeria and Morocco. It often exceeds a height of 60 m , and can Uve for 750 years or more. T w o related species, C. brevifolia and C. libani, occur at the eastern end of the Mediterranean, and C. deodara forms extensive forests in the Himalayas. The surviving forests of C. atlántica in North Africa cover more than 200000 ha, a fragment of their former extent. In Algeria there are extensive forests in the Aurès and smaller forests on other mountains. In Morocco C. atlántica is almost confined to the Rif and the Middle and High Atlas. Its upper limit is well defined and occurs at 2700-2800 m . A t altitudes higher than that the climate is too dry and cold, and Juniperus thurifera becomes dominant. T h e lower limit at which Cedrus forest occurs is less precise and has been obscured by m a n . It is sometimes as low as 1350 m . Isolated trees locally descend to 900 m . Cedrus atlántica is characteristic of the cold humid and subhumid 'étages'. The most luxuriant stands occur on mountain slopes which intercept rain-bearing winds from the Atlantic or Mediterranean, but in parts of its range, as in the High Atlas, the rainfall m a y be as low as 364 m m per year. W h e n , however, it is as low as this it is well distributed and, b y contrast to the general situation in North Africa, an appreciable amount of rain regularly falls in September, and the months of June, July and August, although dry, are not devoid of rain. C . atlántica grows o n a wide range of soils derived from limestone, basalt, schist, marl, granite, and sandstone. At its lower limits C. atlántica is replaced by Juniperus phoenicea, Pinus halepensis, P. pinaster or Quercus ilex. In the wettest regions it is sometimes mixed with deciduous oaks and Abies pinsapo or A . numidica. It is never in contact with Tetraclinis articulata. Although m a n y of the associates of Cedrus atlántica are typical Mediterranean species, Eurosiberian linking species are well represented, especially in the wetter types. Thus in the Rif, Carex leporina, Digitalis purpurea, Luzula fosteri and Solidago virgaurea are plentiful. In the wettest places in marshes and near springs Eurosiberian species are m o r e numerous and include Athyrium filixfemina, Carum verticillatum, Nardus stricta, Parnassia palustris, Pinguicula vulgaris, Primula vulgaris, Sieglingia decumbens, Triglochin palustris and Viola palustris. In
156
Vegetation of the floristic regions
very h u m i d ravines Cedrus is replaced b y Betula péndula, m o s t of w h o s e associates are Eurosiberian species. In addition to those already mentioned are: Alnus glutinosa, Aquilegia vulgaris, Carex distorts, Dryopteris filix-mas, Luzula multiflora, L . sylvatica, Osmunda regalis, Rhamnus frángula (Frángula alnus), Salix cinérea, S. purpurea a n d Sanícula europaea. In the wetter Cedrus forests in the Atlas M o u n t a i n s a n d Algerian coastal ranges C . atlántica is usually accompanied b y Quercus ilex, Acer monspessulanum, Ilex aquifolium, Lonicera arbórea, Sorbus aria, S. torminalis, Juniperus oxycedrus, Hederá helix, Fraxinus xanthoxyloides a n d Taxus baccata. T w o rarer associates are Cytisus battandieri a n d Prunus padus. Eurosiberian species, though less plentiful than in the Rif, are nevertheless prominent. Degradation of these forests gives rise first to thickets of Quercus ilex with Juniperus oxycedrus, Crataegus a n d climbers (Rosa, Rubus, Asparagus), a n d then to Festuca grassland with scattered bushy Q . ilex, J. oxycedrus or Crataegus. Juniperus thurifera is present in the c a n o p y of the drier a n d m o r e continental Cedrus forests, which is always open. This type occurs o n the southern slopes of the M i d d l e Atlas, the eastern, Mediterranean parts of the H i g h Atlas a n d in the Aurès. Quercus ilex is rare. Degradation gives rise to a m i x e d c o m m u n i t y of grasses such as Festuca hystríx, a n d spinous Altimontane shrubs such as Bupleurum spinosum a n d Erinacea anthyllis.
Abies pinsapo and Pi. numidica forest Refs.: Emberger (1939, p. 131-2); Métro (1958, p. 32-4); Peyerimhoff (1941, p. 47); Quézel (1956, p. 18-24). Phots.: Quézel (1956: 3, 4a). Abies pinsapo a n d A . numidica belong to a group of ten closely related species which collectively s h o w a n interrupted circum-Mediterranean distribution. T h e group as a whole is closely related to the central E u r o p e a n A . alba. A . pinsapo is only found in southern Spain a n d a small part of the Rif above C h e c h a o u e n where it occupies 15000 h a o n calcareous soils. T h e M o r o c c a n plant is given separate recognition as subsp. marocana. A . numidica is confined to a few hundred hectares o n the twin summits of B a b o r a n d T a b a b o r in Algeria. Both species occur in the h u m i d 'étage'. A . pinsapo subsp. marocana, which attains a height of 2 0 m a n d a diameter of 1.5 m , b e c o m e s plentiful at 1 5 0 0 - 1 6 0 0 m but descends in ravines to 1 3 0 0 m . It is usually scattered but locally forms dense forests o n inaccessible northern slopes. A . pinsapo forests always include a few Cedrus atlántica, Quercus ilex, Q . fa*ginea, Acer granatense a n d Taxus baccata. T h e floristic composition of the u n d e r w o o d is similar to that of adjacent Cedrus forests. A . numidica also occurs o n calcareous islands inside the m a i n distribution of Cedrus. It is usually m i x e d with Taxus baccata, Q . fa*ginea, Q . afares, Q . ilex, Acer
campestre, A . obtusatum, Sorbus torminalis, S. aria, S. domestica, Populus trémula a n d Ilex aquifolium. Juniperus thurifera forest Refs.: Boudy (1948, p. 135; 1950, p. 754-9); Emberger (1939, p. 86-91); Métro (1958, p. 78); Peyerimhoff (1941, p. 50). Phots.: Boudy (1948: 1; 1950: 118-21); Emberger (1939: 1-2); Métro (1958: 6). Juniperus thurifera has a disjunct distribution in the western Mediterranean. It occurs in the French Alps, the Pyrenees, central Spain a n d the M a g h r e b . In Algeria it is rare a n d is confined to the Aurès, but in M o r o c c o it is m u c h m o r e extensive a n d covers 5 0 0 0 0 ha. It is found o n nearly all M o r o c c a n mountains but is absent from the Rif a n d the western Anti-Atlas. It reaches its m a x i m u m development o n the southern slopes of the G r a n d Atlas. J. thurifera, which is indifferent to substrate, is virtually confined to the cold semi-arid 'étage' between 1800 a n d 3 1 5 0 m . T o w a r d s the lower limits of its range below 2 2 0 0 m it is often mixed with Cedrus atlántica. W h e n unmolested it is a magnificent tree of massive proportions, u p to 15 m tall a n d with a bole 5 m in diameter. M o r e frequently, however, it is mutilated b y h e r d s m e n w h o , in times of s n o w , lop off the branches for fodder a n d fuel. / . thurifera does not regenerate from suckers, and, because of browsing animals, rarely from seed. It resists fire, however, m u c h better than Cedrus, a n d is often the only remnant of m i x e d Cedrus atlántica, Juniperus thurifera forest. Although it n o longer forms forests above the Cedrus-Juniperus zone there is little doubt that it w o u l d d o so if it were not for h u m a n influences. / . thurifera has few faithful associates. M o s t species also occur at higher or lower altitudes than Juniperus itself. A b o v e 2 5 0 0 m notable associates include Bupleurum spinosum, Prunus prostrata a n d Daphne laureola. B u s h y trees often found with it include Crataegus laciniata, Buxus sempervirens a n d Lonicera arbórea. Mediterranean
deciduous forest
Refs.: Boudy (1948, p. 140-1; 1950, p. 252-98); Emberger (1939, p. 132-4); Métro (1958, p. 46-8); Peyerimhoff (1941, p. 52-3, 55); Quézel (1956). Phots.: Boudy (1950: 34-7); Métro (1958: 2); Quézel (1956: 1-2,4b, 5b). There are three species of deciduous o a k in N o r t h Africa, namely Quercus fa*ginea (Q. lusitanica, 'chêne zéen'), Q . pyrenaica (Q. toza, 'chêne tauzin') a n d Q . afares ('chêne afarès'). Quercus fa*ginea extends from Iran to the Iberian peninsula but is absent from Italy, France, a n d the Balearics. It is widespread in N o r t h Africa from M o r o c c o to Tunisia but only in rather small, widely scattered stands. Q . pyrenaica is essentially an Atlantic species, occurring in M o r o c c o , Spain a n d France. Q . afares is confined to Algeria. T h e total area occupied b y deciduous o a k forests in the M a g h r e b is
The Mediterranean regional centre of endemism
small, amounting to n o more than 100000 ha, nearly all in the humid 'étage'. T h e deciduous oak forests of Algeria have been the subject of a profound study by Quezel(1956).
Quercus fa*ginea/oresi Q. fa*ginea is a large tree reaching a height of 30 m and a diameter of 1.5 m . It is deciduous for a few weeks in winter but the dead, brown leaves are retained for several months, especially on young trees and are shed only a few weeks before the n e w ones come, while in s u m m e r its dense crown casts a heavy shade, beneath which it regenerates freely. In Q. fa*ginea forest, the soil is always moist and the underwood usually consists of saplings of the dominant. In Morocco it forms rather small scattered populations, principally in the Rif, the Middle Atlas and in deep valleys o n the El-Harcha-Oulmes plateau. It is confined to the subhumid and humid 'étages' and extends from sea-level in Tanger to 1800 m in the High Atlas. It occurs o n m a n y types of parent material, but the most impressive stands are o n rich volcanic soils. In the Rif there are m a n y islands of Q. fa*ginea forest o n Quaternary volcanic soils in a matrix of Quercus ilex and Cedrus forest on the surrounding calcareous plateau. Quercus fa*ginea usually occurs in pure stands or is diluted by only a few scattered individuals of Q. ilex or Q. suber. Even at the limits of its range it does not mix freely with other species, except w h e n it has been subjected to fire and subsequently invaded by Q. suber. O n calcareous soils, Q. fa*ginea forest has a sparse underwood of Crataegus monogyna, Daphne gnidium, Lonicera etrusca, Rosa and Rubus ulmifolius, but the very dense field layer includes Bromus erectus, Cynosurus echinatus, Festuca triflora, Geum syhaticum, Milium vernale, Pimpinella villosa and several species of Lathyrus, Trifolium and Vicia. O n basalt the underwood, in which Cistus laurifolius, Cytisus battandieri and Viburnum tinus play in important role, is better developed. A t higher altitudes several species equally characteristic of Cedrus forest put in an appearance: Acer monspessulanum, Ilex aquifolium, Sorbus torminalis etc. T h e Q. fa*ginea forests of the Rif differ from the others only in theirrichnessin northern species such as Digitalis purpurea and Prunus avium, and the occurrence of endemics including Linum villarianum and Halimium atlanticum.
Quercus pyrenaica forest Q. pyrenaica forms a series of small populations in the western Rif and Tangier. It occurs pure, or mixed with Q. fa*ginea, Q. ilex or Q. suber. At its upper limits it m a y be mixed with Cedrus. T h e pure forest occurs in a zone where mist is frequent even in summer. In humid ravines ferns are plentiful, especially Aspidium aculeatum, Asple-
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nium adiantum-nigrum, Athyrium filix-femina, Blechnum spicant and Pteridium aquilinum. Quercus afares/wasf This type is similar to Q. fa*ginea forest but occurs o n somewhat drier slopes and at higher altitudes. Q. fa*ginea is often scattered in the canopy. T h e underwood is denser than that of Q. fa*ginea forest but is still sparse compared with that of typical Q. ilex or Q. suber forest. Associates include Ampelodesma mauritanicum, Cytisus villosus, Erica arbórea, Genista tricuspidata, Paeonia atlántica and Ruscus aculeatus. Mediterranean bushland and thicket (mapping units 10, 23, and 78) Refs.: Boudy (1950, p. 436-^tl); Emberger (1939, p. 91-4, 117-18). Syn.: la brousse semi-aride à Olivier-Pistachier-C/iamaeropí (Emberger); Oleo-lentiscetum. It is possible that the climax vegetation occurring on clay soils in the semi-arid 'étage' was bushland or thicket, or, at best, scrub forest dominated by Olea europaea rather than true forest. However, since the original vegetation has virtually disappeared during centuries of intensive cultivation, the matter must remain largely conjectural. T h e degraded remnants of such vegetation are briefly described below. T h e spontaneous form of Olea europaea occurs more or less throughout the Mediterranean Region, but is absent from France. It can grow into a small tree, 10-12 m high, with a short, stout, often twisted, trunk, but is more often seen as 4-5 m high coppice or as small heavily browsed bushes. In the Maghreb it is characteristic of the semi-arid 'étage' and occurs only exceptionally in the subhumid 'étage'. It is virtually absent from the arid and humid 'étages'. Its upper limit in the m o u n tains depends o n humidity. It ascends to only 1200 m in the Algerian Tell, but attains 1650 m in the drier Grand Atlas. If often occurs in forests in the semi-arid 'étage' dominated by Argania spinosa, Juniperus phoenicea, Pinus halepensis, Quercus ilex, Q. suber or Tetraclinis articulata. All these species avoid heavy compact clay soils. T h e latter have been intensively cultivated for centuries and it is difficult to reconstruct their original vegetation. Emberger suggests that it was a bushland or scrub forest dominated by Olea europaea and Pistacia lentiscus with Pistacia atlántica and Chamaerops humilis. Within the area of the Olea-Pistacia community the smallest outcrops of rock or islands of sand support other species, in particular Tetraclinis, Q. ilex, and Q. suber. T h e extensive forests of Q. suber near Rabat which separate the Olea-Pistacia area of the southern Rif from that of Chaouia, illustrate the w a y in which, within the semi-arid 'étage', the vegetation is determined by the soil. M a n y of the associates of Olea and Pistacia lentiscus also occur in Q. suber and Tetraclinis c o m munities in the semi-arid 'étage'.
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Vegetation of the floristic regions
M o s t of the land formerly occupied b y the OlearPistacia c o m m u n i t y , w h e r e it is not cultivated, supports extremely degraded vegetation characterized b y the a b u n d a n c e of the dwarf p a l m Chamaerops humilis, occurring mostly in its acaulescent form. Larger w o o d y associates, which are rare, besides Olea a n d Pistacia lentiscus, include Ceratonia siliqua, Pistacia atlántica, Ziziphus lotus, Crataegus monogyna, Rhamnus oleoides, Rhus pentaphylla a n d Jasminum fruticans. A m o n g smaller plants the following are conspicuous: Anagyris foetida, Asparagus albus, A . stipularis, Calicotome villosa a n d Daphne gnidium. G e o p h y t e s are particularly well represented in the Chamaerops community by species of Aceras, Allium, Anacamptis, Asphodelus, Bellevalia, Colchicum, Crocus, Dipcadi, Erythrostictus, Gagea, Gladiolus, Iris, Leucojum, Muscari, Narcissus, Ophrys, Orchis, Omithogalum, Anthericum, Romulea, Scilla and Urginea. These plants begin their growth several weeks before the first rains appear and so announce the end of summer. In spring the degraded plains and fallow lands of Morocco are like an immense multi-coloured garden or a gigantic Persian or Moroccan carpet because of the profuse flowering of brightly coloured annuals and geophytes. Large patches of orange Calendula algeriensis alternate with or intermingle with purple Fedia, violet Linaria, yellow Diplotaxis and Chrysanthemum, white Ormenis and blue Convolvulus gharbensis, C. tricolor and Echium, a m o n g which emerge the spikes of Gladiolus byzantinus a n d Asphodelus microcarpus, and large clumps of Ferula communis and Foeniculum vulgare. In M o r o c c o the Olea-Pistacia c o m m u n i t y also occurs locally o n clay soils in the s u b h u m i d 'étage' in Tangier. H e r e there are thickets of Pistacia lentiscus a n d Olea, associated with Acanthus, Clematis flammula, Echium boissieri (pomponium), Myrtus communis, Phillyrea angustifolia, Quercus coccifera, Teucrium fruticans a n d the grass Ampelodesma mauritanicum.
Mediterranean shrubland W i t h the exception of halophilous shrubland (page 266) a n d the shrublands of the sub-Mediterranean transition zone, true Mediterranean shrubland is almost confined to high mountains above the timber-line, or is secondary. Altimontane Mediterranean shrubland ( m a p p i n g unit 2 3 ) Refs.: Emberger (1939, p. 138-45); Quézel (1957a, p . 109-78, 193-211,418-20); Taton (1966); White ( M S , 1974). Phots.: Emberger (1939: 9.1-4); Quézel (1957a: 5, 6, 8, 14, 20). Syn.: les garrigues montagnardes à xérophytes épineux (Quézel); l'horizon à xérophytes épineux en coussinets (Emberger). D w a r f shrubland dominated b y dense, cushion-shaped, very spinous shrubs is o n e of the m o s t widespread a n d
characteristic vegetation types o n the N o r t h African mountains. It grows under a typical Mediterranean climate, since the rain falls during the cold season as in the plains. It is, however, a n extreme climate and s n o w lies for several m o n t h s in winter. T h e soils frequently s h o w polygonal structure d u e to frost action a n d Quézel regards t h e m as periglacial. Altimontane shrubland nearly always grows o n skeletal soils a n d during the s u m m e r is exposed to very dry conditions. In M o r o c c o , Altimontane shrubland represents the climatic climax between the tree-line a n d 3 8 0 0 - 3 9 0 0 m . It also occurs very extensively inside the forest belt d o w n to c. 2 0 0 0 m o n the drier m o u n t a i n slopes; in this situation it is mostly secondary. There c a n b e little doubt, however, that, before the forests were destroyed, the shrubland species formed small colonies o n shallow soil a n d rocky outcrops w h e r e the trees were unable to form a closed canopy. Because of the widespread destruction of forest it is difficult to determine the lower climatic limit of Altimontane shrubland. F o r M o r o c c o , E m b e r g e r believes it to occur at about 2 8 0 0 m . Altimontane shrubland is extensively developed in M o r o c c o , especially in the Great Atlas a n d o n the s u m m i t s of the M i d d l e Atlas. In the Anti-Atlas it is only found o n the massif of Siroua. T h e Rif is too l o w for its occurrence. In Algeria it is m u c h m o r e restricted than in M o r o c c o , but it occurs o n the s u m m i t s of the Djurdjura a n d the Aurès. Physiognomically similar vegetation occurs o n all the high mountains in the Mediterranean basin, but floristically similar types are found only in southern and eastern Spain. T h e flora of Altimontane shrubland is poor and uniform, but rich in endemic species. According to Quézel, the total Altimontane flora in the M a g h r e b a m o u n t s to n o m o r e than 6 5 0 species. M o s t of t h e m are confined to the Mediterranean Region. O n e hundred a n d sixty species are endemic to the high mountains of the M a g h r e b . Nearly all belong to Mediterranean endemic genera. Eurosiberian species are relatively poorly represented, a n d Carex capillaris is o n e of the few northern species which in the M a g h r e b is confined to the high mountains. T h e dominants of Altimontane shrubland are dwarf shrubs. In M o r o c c o there are 18 species, n a m e l y Alyssum spinosum (3850 m ) , Amelanchier ovalis (3500 m ) , Arenaria dyris (3750 m ) , A . pungens (3790 m ) , Berberís hispánica (3200 m ) , Bupleurum spinosum (3400 m ) , Cytisus balansae (3600 m ) , Erinacea anthyllis (3600 m ) , Juniperus communis (3300 m ) , J. oxycedrus (3150 m ) , Lonicera pyrenaica (3500 m ) , Ononis atlántica (3250 m ) , Prunus prostrata (3200 m ) , Rhamus alpinus (3200 m ) , Ribes alpinum (3300 m ) , R . uva-crispa (3400 m ) , Sorbus aria (3000 m ) a n d Vella mairei (3200 m ) . U p p e r altitudinal limits are s h o w n in brackets. T h e spinous cushion plants (Alyssum, Arenaria pungens, Bupleurum, Cytisus a n d Erinacea) are normally about 0.5 m tall. Because of heavy grazing the ground between t h e m is often almost bare, a n d other species in the c o m m u n i t y can only g r o w with their protection.
The Mediterranean regional centre ofendemism
T h e spinous cushion plants are rare or absent from certain habitats such as escarpments, mobile screes, earthy slopes, and from the highest altitudes, where they are replaced by herbaceous communities.
Secondary Mediterranean shrubland (maquis and garrigue) (mapping units 10,23, and 78) Refs.: Gimmingham & Walton (1954); Ionesco & Sauvage (1962); TomaseU (1976). Phots.: Gimmingham & Walton (1954: 1, 2,4-8). T h e climate of Mediterranean Africa is a forest climate, and forest, scrub forest, or, locally, bushland and thicket represent the climax nearly everywhere below the natural tree line. A s one approaches the Sahara there is a progressive diminution in the height of the vegetation, and various types of bushland and shrubland occur in the Mediterranean-Sahara transition zone (Chapter XVIII). Within the Mediterranean Region proper, nearly all communities dominated by shrubs have been derived from forest, scrub forest, or bushland. S o m e of the taller shrublands are sometimes referred to as 'maquis' or 'macchia', and some of the shorter as 'garrigue', but considerable confusion is associated with the application of these terms. Classical maquis is dense and tall (up to c. 4 m ) , difficult to penetrate, and is dominated by Erica arbórea and Arbutus unedo. It occurs on siliceous soils and degrades to a grassland dominated by Ampelodesma mauritanicum. T h e term garrigue is derived from the Catalan n a m e for Quercus coccifera (garric) and was originally applied to low bushy communities dominated by that species which occur on calcareous soil. Its application is sometimes extended to embrace all open shrubby c o m munities of m e d i u m height occurring on calcareous soil in the Mediterranean Region. T h e value of this distinction is doubtful and in practice m a n y authors have used both terms in senses different from those quoted above. T h e importance of the lithological origin of the soil in the Maghreb has probably been exaggerated, since m a n y species, including Arbutus unedo and Quercus coccifera, grow equally well o n calcareous and siliceous soil. In North Africa, Q. coccifera has a restricted distribution. Its most characteristic habitat is Olea europaea, Pistacia lentiscus, Ceratonia siliqua bushland on clay soils. It is also frequent in maquis and is not found on skeletal calcareous soils (P. J. Stewart, pers. c o m m . ) . In practice, the terms maquis and garrigue have usually been defined in the past in such a w a y as to cover only a part of the wide range of secondary shrublands that occur in the Mediterranean. S o m e authors, however, e.g. TomaseUi (1976) in a review of maquis covering the whole of the Mediterranean basin, take a
159
more comprehensive view and draw a purely arbitrary distinction based on height. T o TomaseUi, maquis is a formation more than 2 m tall; it is usually dense and composed of w o o d y plants without a well-defined trunk. It is indifferent to substrate. Since the stature and density of secondary shrubland depends as m u c h on the intensity of degradation as on the nature of the parent material, and floristic composition varies in a complex manner from place to place in relation to the floristic composition of the original forest, there seems to be little justification for the use of the terms maquis and garrigue in a pan-African classification, though doubtless endless litigation will continue to be associated with their local use. B y contrast, the use of a vernacular term, 'fynbos' (page 132), to designate a somewhat physiognomically similar type of shrubland in the Cape Region, is free from ambiguity, possibly because the vegetation it refers to is a regional climatic climax which has considerable floristic uniformity throughout its range. The structure andfloristiccomposition of secondary Mediterranean shrubland are so variable that little would be gained from an attempt to describe them. S o m e examples are briefly described elsewhere in this chapter, along with the forests from which they have been derived. G i m m i n g h a m & Walton (1954) briefly describe three stages in the degradation of Cupressus sempervirens, Juniperus phoenicea, Olea europea, Quercus coccifera, Ceratonia siliqua scrub forest on calcareous soil in Cyrenaica. The first stage is dominated by Arbutus pavarii often associated with Ceratonia siliqua, Phillyrea angustifolia, and Pistacia lentiscus, with an irregular field layer of Poterium (Sarcopoterium) spinosum, Cistus parviflorus etc. Arbutus pavarii, which forms a thicket of slender stems 1.8-3 m high, is less susceptible to grazing than saplings of trees which have a single trunk, and so persists under moderate grazing. It is, however, hable to perish under conditions of more heavy grazing, w h e n Pistacia lentiscus forms dense thickets 1.5-2.4 m high. Under the most intensive grazing even Pistacia disappears and its place as dominant is taken by the low shrub Poterium spinosum. In the eastern Mediterranean, communities dominated by Poterium are referred to as 'batha'.
Mediterranean anthropic landscapes (mapping unit 78) T h e most fertile lowlands have been cultivated since R o m a n times, and few vestiges of the natural vegetation remain. W h e a t is the most widely planted crop, but peas, beans, and onions are plentiful, and there are m a n y groves of olive, Citrus, fig, and Vitis. T h e hedges are mostly of Agave, Acacia karroo, Arundo donax and Opuntia. T h e latter is frequently
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Vegetation of the floristic regions
naturalized. Eucalyptus and Pinus halepensis are planted locally as wind-breaks and for fuel and timber, but in general the landscape is treeless. The sparse plant cover of shallower soils is grazed by sheep and cattle, but owing to overstocking there is
m u c h soil erosion and in places the stony mantle supports only Ziziphus lotus, dwarf Chamaerops, and unpalatable herbs such as Ferula communis, Asphodelus microcarpus and Urginea marítima. Marshy hollows are often dominated by Juncus acutus.
viii/ix T h e Afromontane archipelago-like regional centre of endemism and the Afroalpine archipelago-like region of extremefloristicimpoverishment Geographical position and area
Geographical position and area
Geology and physiography
T h e Afromontane Region is an archipelago-like centre of endemism (White, 1978a) which extends from the L o m a M t s and the Tingi Hills ( 1 1 ° W . ) in Sierra Leone in the west to the Ahl Mescat M t s (49° E.) in Somalia in the east, and from the R e d Sea Hills (17° N . ) in the Sudan Republic in the north to the C a p e Peninsula (34° S.) in the south. A few Afromontane species descend almost to sea-level even in the tropics, but outside the Afromontane Region they are always very rare. In the tropics most Afromontane communities are found only above 2000 m , but where the climate is more oceanic, as in the West U s a m b a r a M t s in Tanzania, they can occur as low as 1200 m . Further south, where latitude compensates for altitude, they descend progressively further, and in the C a p e Region exclaves of Afromontane forest are found only a few hundred metres above sea-level. T h e inclusion of the West African mountains west of C a m e r o u n , and the highlands of Angola, in the Afromontane Region is still problematical, since the Afromontane species occurring there appear to be diluted by the presence of m a n y lowland species. (Area: 715000 k m 2 . )
Climate Flora Mapping units Vegetation Afromontane forest Afromontane rain forest Undifferentiated Afromontane forest Single-dominant Afromontane forest Juníperas procera forest Widdringtonia cupressoides forest Hagenia abyssinica forest Dry transitional montane forest Afromontane bamboo Afromontane evergreen bushland and thicket Afromontane and Afroalpine shrubland Afromontane and Afroalpine grassland Mixed Afroalpine communities In tropical Africa In South Africa
Geology and physiography T h e Afromontane 'archipelago' is very diverse in lithology and physiography, which have been little studied from a botanical point of view. S o m e of the largest 'islands' and m a n y of the smaller ones are largely volcanic in origin, though the lavas are of different ages. M o s t of the Ethiopian highlands are formed of basalt, though Precambrian rocks outcrop locally. T h e oldest volcanic rocks date from the Eocene and have been intermittently augmented by subsequent eruptions which have continued into the Quaternary. T h e K e n y a highlands are mostly formed of volcanic deposits, including phonolite, nephelinite and basalt, which were erupted in post-Miocene time during the formation of the eastern Rift Valley. B y contrast, the Cherangani Hills (3600 m ) are composed of Precambrian metamorphic rocks with conspicuous quartzite ridges.
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Vegetation of the floristic regions
The highest parts of the Natal Drakensberg and adjacent Lesotho are capped by more or less horizontal basaltic lava flows which terminate the Stormberg Series of Triassic age. Adjacent less-elevated parts of the Afromontane Region, however, are underlain by Karoo sediments of the Stormberg and Beaufort series. The Kivu ridge and the contiguous uplands which extend from the northern end of Lake Tanganyika to Ruwenzori are largely composed of Precambrian rocks, but with local islands of volcanic deposits including those formed by the still active Virunga volcanoes (4507 m ) . The Cameroun Highlands are formed partly of volcanic, partly of ancient crystalline rocks. M o u n t Cameroun (4095 m ) , a still active volcano, stands separate from the main range. O f the more isolated mountains some are volcanic in origin, e.g. M t Elgon (4315 m ) , M t Meru (4566 m ) and M t Kilimanjaro (5 890 m ) , whereas others are formed of crystalline basem*nt rocks. S o m e of the latter, e.g. the Chimanimani Mts in Zimbabwe, represent the resistant remnants of the uplifted rim of the Great African Plateau, while others, notably Ruwenzori (5119 m ) , have been uplifted by compressional forces associated with rift faulting.
Climate T h e climate is extremely varied but reliable published records are sparse. Information o n the Afroalpine zone is provided b y Hedberg (1964), o n the Austro-afroalpine zone b y Killick (1978a, 19786, 1978c), o n M t C a m e r o u n by Richards (1963¿>), o n the forest belt in Malawi b y C h a p m a n & White (1970), and o n the mountains at the eastern end of the Zaire basin b y Bultot (1950, 1971-74) and Scaëtta (1933,1934). In the forest belt, m e a n annual rainfall is usually m o r e than 1000 m m , but is less in drier types transitional to lowland vegetation. A b o v e the forest belt precipitation diminishes and in the Afroalpine belt of s o m e mountains appears to be m u c h less than 1000 m m per year. Cloud is a feature of most mountains but its importance is very uneven and it has been little studied. T h e incidence of frost varies considerably from complete absence o n s o m e lower slopes to a nightly occurrence o n the highest summits. (See Fig. 14.)
Flora There are at least 4 0 0 0 species of which c. 3000 are endemic or almost so. Endemic and near-endemic families. Barbeyaceae, Oliniaceae. Curtisia (Cornaceae) is sometimes given family rank. Barbeya, which also occurs in the Y e m e n , is not strictly Afromontane, but is m o r e characteristic of the ecotone between dry Afromontane forest and Somalia-Masai evergreen bushland.
Endemic genera. A b o u t one-fifth of the tree genera, including Afrocrania, Balthasaria, Ficalhoa, Hagenia, Kiggelaria, Leucosidea, Platypterocarpus, Trichocladus, and Xymalos, are endemic. For smaller plants the proportion is probably less and includes Ardisiandra, Cincinnobotrys, and Stapfiella. Linking elements. See White (1978a, p . 475-80).
Mapping units 19a. Undifferentiated Afromontane vegetation. 65. Tropical altimontane vegetation. 66. South African altimontane vegetation. In addition to the above, Afromontane species also occur in the following mapping units: 4. Transitional rain forest. 13. The Fouta Djalon mosaic of lowland rain forest and secondary grassland with montane elements. 17. Cultivation and secondary grassland replacing upland montane forest. 19b. Undifferentiated Sahelomontane vegetation. 32. Jos Plateau mosaic. 33. M a n d a r a Plateau mosaic.
Vegetation O n any particular mountain there is usually a very wide range of vegetation. Extreme types m a y have few species in c o m m o n , but all types are intimately connected b y complex series of intermediates. T h efloristicdifferences between extremes o n a single mountain are usually greater than the differences between the Afromontane assemblage as a whole o n that mountain and the assemblage found o n nearby, or indeed o n distant, mountains so that the collective flora of the archipelago shows a remarkable continuity and uniformity. O n most mountains the lowermost vegetation is forest, beneath which one would expect to find a transition zone connecting the Afromontane and lowland phytochoria. Nearly everywhere, however, the vegetation of this transition zone has been destroyed b y fire a n d cultivation, but the remnants of a dry type of transition forest are briefly described below. Forests transitional to Guineo-Congolian, Zanzibar-Inhambane and Tongaland-Pondoland communities are described in Chapters I, XIII, and X V respectively. O n nearly all African mountains the vegetation diminishes in stature from the lower slopes to the summit, but this regularity is so often modified b y local features of aspect, exposure, incidence of frost, and depth of soil, and b y overall patterns of climate dependent o n the size and configuration of the mountain in relation to distance from the sea and other sources of moisture, that generalized schemes of zonation, even for relatively restricted regions, are impossible to devise. Nevertheless, the three broad belts,
The Afromontane archipelago-like regional centre of endemism and the Afroalpine archipelago-like region
ADDIS ABEBA(2440m) 15-9* 1302 (23-28) ^
ADI-UGRI (2022m)l93°730 ( 9 )
FIG. 14. Climate and topography of the Afromontane archipelago-like regional centre of endemism (VIII) and the Afroalpine archipelago-like region of extreme floristic impoverishment (IX) (The two phytochoria, shown as solid black areas, are not separately distinguished)
163
164
Vegetation of the floristic regions
forest, Ericaceous and Afroalpine, recognized by Hedberg (1951) for the high mountains of East Africa, can usually be detected, but the details of zonal replacement vary greatly, even on different slopes of the same mountain. In the C a p e Region, Afromontane vegetation, which is represented there only b y forest, is n o longer associated with the highest parts of mountains but is confined to the lower slopes. Except o n the wettest mountains where the original vegetation is sometimes well preserved the most widespread vegetation is secondary, fire-maintained grassland. T h e Ericaceous and Afroalpine belts o n the high mountains of tropical East Africa are m a p p e d collectively as Altimontane vegetation (unit 65). Ecologically similar butfloristicallysomewhat different vegetation in South Africa is m a p p e d as unit 66. N o attempt is m a d e to m a p the remaining Afromontane types separately, except that large areas of Juniperus forest and mixed Afromontane forest are indicated by letters. T h e small areas shown as Afromontane in Angola and in the L o m a - M a n highlands in West Africa harbour m a n y Afromontane species which d o not seem, however, to form extensive Afromontane communities. Afromontane forest M o s t Afromontane tree species have wide geographical distributions and wide ecological amplitudes. M a n y also exhibit a wide range of growth forms. For these reasons classification of the forests they form is difficult, and the latter are probably best regarded as comprising a continuum which is only slightly structured. It is, however, convenient to separate off the most luxuriant types as Afromontane rain forest, and to give individual treatment to certain extreme floristic variants dominated by single species, although it appears that the latter are nearly always secondary in origin. Rainfall received b y Afromontane forest varies from 800 m m to considerably more than 2500 m m per year. Nevertheless, the distinction drawn by earlier workers (e.g. Greenway, 1973) between 'wet' and 'dry' types is difficult to apply, chiefly owing to the wide tolerance of m a n y dominant species. Little has been published o n the transition from Afromontane forest to lowland communities. D r y transitional Afromontane forest in East Africa is described in this chapter, and the bushland which replaces it at lower altitudes is dealt with in Chapter IV. Transitional rain forest is described in Chapters I, XII, and XIII. Afromontane rain forest (mapping unit 19a) Refs.: Chapman & White (1970); Langdale-Brown, Osmaston & Wilson (1964, p. 42-3, 109-10, p.p.); Lewalle (1972, p. 107-14); Mildbraed (1914, p. 623-6); Pichi-Sermolli (1957, p. 82-4); Pitt-Schenkel (1938); Pócs (19766, p. 4867); White (1978a, p. 485; M S , 1952,1975,1976).
Phots.: Chapman & White (1970: 1-7, 12, 24, 40-4); LangdaleBrown et al. (1964: 3). Profiles: Chapman & White (1970: 1-4, 6); Lewalle (1972: 21, 23). Syn.: foresta umida sempreverde montane (Pichi-Sermolli, 1957); forêt ombrophile de montagne: horizons inférieur et moyen (Lewalle, 1972, p.p.); Pygeum moist montane forest (Langdale-Brown et al., 1964); submontane rain forest (Pócs, 19766); submontane seasonal rain forest (White, in Chapman & White, 1970). Afromontane rain forest is very similar in structure and physiognomy to certain types of Guineo-Congolian lowland rain forest. A t the species level, however, it is almost completely different, but m a n y of its species are closely related to Guineo-Congolian species, or have their closest relatives elsewhere in the lowland tropics. These are the 'nephews' and 'orphans' of White (in C h a p m a n & White, 1970). The most characteristic tree species of Afromontane rain forest include Aningeria adolfi-friedericii, Chrysophyllum gorungosanum, Cola greenwayi, Cylicomorpha parviflora, Diospyros abyssinica, Drypetes gerrardii, Entandrophragma excelsum, Ficalhoa laurifolia, Mitragyna rubrostipulata, Myrianthus holstii, Ochna holstii, Ocotea usambarensis, Olea capensis, Parinari excelsa, Podocarpus latifolius (including milanjianus), Prunus africana, Strombosia scheffleri, Syzygium guiñéense subsp. afromontanum, Tabernaemontana johnstonii, and Xymalos monospora. Afromontane rain forest occurs mostly between 1200 and 2500 m , but its precise altitudinal limits vary greatly according to distance from the equator, proximity to the sea, and size and configuration of the massif o n which it occurs. It is found on the wetter slopes of most of the higher mountain massifs from southern Ethiopia to Malawi. T h e m e a n annual rainfall of Afromontane rain forest lies mostly between 1250 and 2500 m m , but is sometimes higher. There is usually a dry season lasting from one to five months, but dry-season mists are frequent. This m a y explain the fact that upland rain forest is often m u c h less deciduous than lowland semievergreen rain forest experiencing a similar rainfall. Apart from secondary species, only a few of the larger tree species, e.g. Aningeria adolfi-friedericii and Entandrophragma excelsum, lose their leaves, and then only for a few days. Frosts occur occasionally but are not severe. Detailed published information on Afromontane forest is available only for Burundi (Lewalle), the West Usambara Mts (Pitt-Schenkel), and Malawi (Chapman & White). In physiognomy Afromontane rain forest is similar to certain types of Guineo-Gongolian lowland rain forest. The trees of the upper stratum are 25-45 m tall (average 30-38 m ) . Their crowns, which are not in lateral contact, are raised well above the middle tree stratum and are heavy-branched and wide-spreading. The middle tree stratum is 14-30 m tall; its crowns are often
The Afromontane archipelago-like regional centre of endemism and the Afroalpine archipelago-like region
narrow and conical and m a y be discontinuous to continuous but do not form a dense canopy. T h e lower tree stratum is 6-15 m tall and usually forms a dense canopy. T h e shrub layer, 3 - 6 m tall, is poorly differentiated from the lower tree layer. T h e herb layer is usually sparse and consists largely of forest grasses and ferns. Lianes and strangling epiphytes are abundant. A m o n g vascular epiphytes, ferns and lycopods are m o r e or less abundant throughout, and species of Begonia, Impatiens, Streptocarpus and Peperomia are widespread. Orchids, though present, are not abundant, and Rhipsalis occurs locally. Epiphytic bryophytes are generally present but are not abundant except in some of the wetter types. Epiphytic lichens are not generally abundant except in the crowns of certain species. A few of the larger trees are buttressed and some are briefly deciduous, but the general impression throughout the year is that the forests are evergreen. Physiognomically, Afromontane rain forest differs from most types of Guineo-Congolian rain forest chiefly in the occurrence of tree ferns (Cyathea) and conifers {Podocarpus). T h e latter, however, are more characteristic of other types of Afromontane forest. There is also a greater degree of bud protection, and 'drip tips' of the leaves are less well developed. Undifferentiated Afromontane forest (mapping unit 19a) Refs.: Aco*cks (1975, p. 18-23, 25-7, 82-6); Chapman & White (1970, p. 107-9, 131-2; 137-9, 148-51); Edwards (1967, p. 174-80); Jackson (1956, p. 365-70); Killick (1963, p. 4357); Letouzey (1968a, p. 325-48); Lewalle (1972, p. 11423); Lind & Morrison (1974, p. 32-43, p.p.); Pichi-Sermolli (1957, p. 73-84, p.p.); Pócs (19766): Richards (19636); von Breitenbach (1972); White (1978a); Wild & Barbosa (1967, p. 10-11). Phots.: Aco*cks (1975: 8); Chapman & White (1970: 45, 46); Dyer (1937: 17); Edwards (1967: 111-15); Killick (1963: 16); Moll (1966: 1-3; 1968c: 10); Philipps (1931); Phipps & Goodier (1962: 2); Richards (19636: 1, 2); von Breitenbach (1972: on pp. 19-26). Profiles: Boughey (1961: 3, 4); Chapman & White (1970: 5); Jackson (1956: 4); Moll (1968c: 6, 7); Richards (19636: 1). Syn.: broad-leaved montane forest (Chapman & White, 1970); Dohne sourveld (Aco*cks, 1974); forêt ombrophile de montagne: horizon supérieur (Lewalle, 1972); highland sourveld (Aco*cks, 1975); Knysna forest (Aco*cks, 1975); mist-belt mixed Podocarpus forest (Edwards, 1967); moist broad-leaved montane forest (Wild & Barbosa, 1967); mountain Podocarpus forest (Edwards, 1967); Natal mist belt 'ngongoni' veld (Aco*cks, 1975); north-east mountain sourveld (Aco*cks, 1975); 'ngongoni' veld (Aco*cks, 1975); Pondoland coastal plateau sourveld (Aco*cks, 1975, p.p.). Undifferentiated Afromontane forest is usually shorter than Afromontane rain forest and despite some floristic overlap is of distinctive composition. It usually replaces rain forest at higher altitudes on the wetter slopes and at comparable altitudes o n the drier slopes. O n some mountains it also occurs below Afromontane rain forest. It usually but not always receives a lower rainfall.
165
M o s t stands of undifferentiated Afromontane forest are floristically mixed, but sometimes after fire they are replaced b y almost pure stands of Juníperas procera, Widdringtonia cupressoides (whytei) or Hagenia abyssinica (see below). T h e majority of tree species in this type are very widespread. A m o n g them are Apodytes dimidiata, Halleria lucida, Ilex mitis, Kiggelaria africana, Nuxia congesta, N. floribunda, Ocotea bullata (including O. kenyensis), Podocarpus falcatus (including gracilior), P. latifolius, Prunus africana, Rapanea melanophloeos s.l. and Xymalos monospora. Indeed this assemblage of species could almost be used to define the Afromontane Region as a whole. N o single species occurs throughout, but the assemblage is represented o n virtually every 'island' of Afromontane vegetation, usually by several species. A H are widespread in South Africa and all except four {Kiggelaria, Nuxia floribunda, Podocarpus latifolius and Xymalos) extend as far north as Ethiopia. Only five {Halleria, Kiggelaria, Nuxia floribunda, Ocotea and Podocarpus falcatus) are absent from West Africa. S o m e species such as Combretum kraussii, Cryptocarya latifolia, C. woodii, Curtisia dentata {fa*ginea), Chionanthus foveolatus, Ptaeroxylon obliquum, Schefflera umbellifera, Scolopia mundii and Podocarpus henkelii (including ensiculus), which are important in South Africa, either d o not cross the L i m p o p o River or are very localized further north. Single-dominant Afromontane forest (mapping unit 19a) Juniperus procera forest Refs.: Chapman & White (1970, p. 108-9, 130-1); Hemming (1966, p. 218-21); Langdale-Brown, Osmaston & Wilson (1964, p. 43-4, 110-11); Lind & Morrison (1974, p. 41-2); Pichi-Sermolli (1957, p. 75-80); White (MS, 1975, 1979); Wimbush(1937). Phots.: Chapman & White (1970: 13-16); Herlocker & Dirschl (1972: 22); Pichi-Sermolli (1957: 11); Wimbush (1937: between pp. 50 and 51). Juniperus procera has a scattered distribution o n the eastern side of Africa from the R e d Sea Hills in the Sudan, Eritrea, and Arabia to the Nyika Plateau in northern Malawi. A single individual has also been reported from the van Niekerk ruins in Z i m b a b w e 1280 k m further south. It mostly occurs on the drier slopes of mountains between 1800 and 2900 m but it occasionally descends to as low as 1000 m . Rainfall is between 1000 and 1150 m m per year, but welldeveloped stands of forest more than 30 m tall occasionally occur where the rainfall exceeds 1250 m m per year (Glover & T r u m p , 1970). It is also present as an emergent in scrub forest and evergreen bushland where the annual rainfall is as low as 650 m m ( H e m m i n g , 1966), and this m a y represent its original habitat. Juniperus procera most frequently forms forests in which it is b y far the most abundant species. It is,
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Vegetation of thefloristicregions
however, a strong hght-demander and does not regenerate in its o w n shade. Its seedlings also seem to be intolerant of a deep layer of h u m u s on the surface of the soil (Gardner, 1926). It is clear that its presence as a forest tree is largely dependent on fire, either natural or m a n - m a d e (Wimbush, 1937).
Widdringtonia cupressoides forest Refs.: Chapman & White (1970, p. 108-9, 162-9); Van der Schijff & Schoonraad (1971, p. 472); White (MS, 1973); Wild & Barbosa (1967, p. 11). Phots.: Chapman & White (1970: 51, 53-7). Profile: Chapman & White (1970: 7). Syn.: dry montane conifer forest (Wild & Barbosa, 1967: 11, p.p.). Widdringtonia cupressoides extends along the eastern side of Africa from Table Mountain in the south to M t Mulanje in the north. Over the greater part of its range it occurs as a small, usually bushy, often multiplestemmed, tree from 4 to 9 m tall, and scarcely ever forms forests. It is only on M t Mulanje that it occurs as a tall forest tree. Although individual trees attain a height of 40 m the main canopy of mature forest is normally c. 27 m high. T h e forests, which occur between 1525 and 2135 m , are very susceptible to fire and the behaviour of Widdringtonia cupressoides in relation to fire is similar to that of Juniperus procera.
Hagenia abyssinica forest Refs.: Chapman & White (1970, p. 131-2); Demaret (1958); Jackson (1956, p. 361); Langdale-Brown, Osmaston & Wilson (1964, p. 43, 110); Lebrun (1942, p. 52-6); Lind & Morrison (1974, p. 48-50); Robyns (1948a, p. xli-xlii); Spinage (1972, p. 198); White (MS, 1975, 1978). Phots.: Lebrun (1942: 23a); Lind & Morrison (1974: 12); Robyns (1937: 3b); Spinage (1972: 7). Profile: Jackson (1956: 2). Syn.: Hagenia woodland (Lind & Morrison, 1974); HageniaRapanea moist montane forest (Langdale-Brown et al., 1964); la forêt-prairie à Hagenia abyssinica (Lebrun, 1942). Hagenia abyssinica is found o n most of the higher mountains between Ethiopia and the Nyika Plateau in northern Malawi. It occurs both o n the wetter mountains such as Ruwenzori, where it is rare, and the drier mountains such as M t M e r u . Its abundance does not seem to b e at all closely related to moisture conditions. Its altitudinal range lies between 1800 and 3400 m but it is normally absent from Afromontane rain forest and the taller types of undifferentiated montane forest. Characteristically Hagenia forms almost pure stands 9-15 m tall in a narrow zone (often interrupted) between taller types of montane forest and the thickets and shrublands of the Ericaceous belt. T h e biggest trees have boles up to 2 m long and 1.6 m in diameter which support massive spreading branches. T h e best-developed stands are clearly forest, though of a simpler structure
than most African forests. Other stands have more the structure of woodland or scrub forest. Hagenia is a heliophyte which can withstand at least some burning, though it is killed by repeated fierce fires. At lower altitudes it is always serai. Thus, in the Imatong M t s w h e n secondary grassland derived from lower montane forest is protected from fire, Hagenia rapidly invades, followed b y other forest species (Jackson, 1956). At higher altitudes, where Hagenia is m u c h more abundant, its status is still uncertain. Lind & Morrison (1974) suggest that Hagenia forest m a y be climax where low night temperatures exclude m a n y other trees, and competition is low. There is little doubt, however, that, even at high altitudes, its abundance is at least partly due to disturbance. O n the Nyika Plateau ( C h a p m a n & White, 1970) it occurs in short broad-leaved montane forest 8-15 m tall, in association with about 20 other tree species. Because the canopy is low and is kept open by large m a m m a l s , heliophilous species like Hagenia can establish themselves. Hagenia, however, is more abundant at the forest edge. Elsewhere, small patches of unmodified forest are surrounded by wide aureoles of forest in the process of degradation b y fire, in which Hagenia abyssinica is usually the dominant species. Repeated burning leads to the replacement of the trees by thicket and ultimately secondary grassland. O n the north-eastern slopes of M t K e n y a (White, M S , 1975) the patches of Hagenia forest just below the Ericaceous belt have been m u c h influenced by buffalo. T h e understorey has been degraded by browsing, and largely replaced by a sward of grass and Trifolium. N o regeneration of Hagenia is to be seen. Lower d o w n the mountain Hagenia forms small groves inside Juniperus forest. In the Virunga M t s , at c. 3000 m , Hagenia, in association with Hypericum revolutum, forms a low forest, 10-12 m high, but there is no regeneration (P. B a m p s , pers. c o m m . ) .
Dry transitional montane forest (mapping unit 19a) Refs.: White (MS, 1973,1975, 1979). T h e drier lower slopes of those East African mountains and uplands which rise from the bushland-covered Somalia-Masai plains formerly supported a dry type of forest in which Afromontane and non-Afromontane species occurred together. Only small fragments remain and there is little published information. There are some well-preserved examples between 1650 and 1800 m near Nairobi where the rainfall is c. 800 m m per year. T h e main canopy is at 15-18 m with émergents up to 25 m high. T h e larger trees include Albizia gummifera (near streams), Apodytes dimidiata, Brachylaena discolor, Calodendrum capense, Cassipowea congoensis (including malosana), Chaetacme aristata,
The Afromontane archipelago-like regional centre ofendemism and the Afroalpine archipelago-like region
Chrysophyllum viridifolium, Crotón megalocarpus, Diospyros abyssinica, Drypetes gerrardii, Euclea divinorum, fa*garopsis angolensis, Manilkara obovata, Markhamia hildebrandtii, Newtonia buchananii (near streams), Olea africana, Phyllanthus discoideas, Schrebera alata, Strychnos usambarensis, Suregada procera, Teclea spp. and other Rutaceae, Trichocladus ellipticus, Uvariodendron anisatum and Warbwgia salutaris (ugandensis).
Afromontane b a m b o o (mapping unit 19a) Refs.: Aco*cks (1975, p. 97); Chapman & White (1970, p. 166); Demaret (1958, p. 332); Fries & Fries (1948, p. 31-9); Glover & Trump (1970, p. 17-21); Greenway (1965, p. 98); Hedberg (1951); Hendrickx (1944, p . 5; 1946, p . 39); Jackson (1956, p. 368, 370); Keay (1955, p. 142); Kerfoot (1964a, p. 298); Langdale-Brown et al, (1964, p. 44, 111); Lebrun (1942, p. 47-9; 1960, p. 89); Letouzey (1968a, p. 336, 338); Lewalle (1972, p. 124-31); Lind & Morrison (1974, p. 45-7); Mabberley (1975a, p. 4); Pichi-Sermolli (1957, p. 84-6); Pócs (1976a, p. 489; 19766, p. 169); Robyns (1937, p. 12-14; 1948, p. xli, xlvii); Snowden (1953, p. 63-4); Spinage (1972, p. 198); Tweedie (1976, p. 240); White ( M S , 1949, 1963, 1973, 1975). Phots.: Langdale-Brown et al. (1964: 7); Lind & Morrison (1974: 10, 11); Robyns (1937: 4a). Profile: Lewalle (1972: 26). Syn.: Arundinaria alpina forest or thicket (bamboo) (LangdaleBrown et al., 1964); foresta a bambú (Arundinaria) (PichiSermolli, 1957); moist bamboo grass thicket (Greenway, 1973, p. 64). Arundinaria alpina occurs o n most of the high mountains in East Africa, from Ethiopia to the Southern Highlands in Tanzania. Further south it is only k n o w n from the North Viphya Plateau, D e d z a M t and M t Mulanje in southern Malawi. In south Africa it is replaced b y A. tessellata. In W e s t Africa it is found sporadically o n s o m e of the mountains of C a m e r o u n , though not o n M t C a m e r o u n itself. T h e extent and vigour of A. alpina varies greatly o n different mountains and the reasons for this are not yet understood. In East Africa A. alpina is mostly found between 2 3 8 0 and 3000 m , but o n M t K e n y a it ascends to 3200 m , and in the Uluguru M t s it descends to 1630 m . It appears to grow most vigorously and to form continuous stands o n deep volcanic soils o n gentle slopes where the rainfall exceeds 1250 m m per year. T h e largest areas are o n the Aberdare R a n g e (65000 hectares, 250* miles), M a u R a n g e (51000 hectares, 200 J miles) and M t K e n y a (39000 hectares, 150 2 miles). O n M t Elgon it has a patchy distribution o n the drier eastern side but forms continuous stretches o n the moister western side. O n Ruwenzori it is poorly developed o n very steep slopes but elsewhere is dominant between 2200 and 3200 m . Arundinaria is almost absent from the 'dry' mountain Kilimanjaro but forms a belt between 2 1 3 0 and 2740 m o n the adjacent and n o less dry M t M e r u (Greenway, 1965). It is mostly of sporadic occurrence in the Cherangani and Uluguru M t s but in the
latter forms pure stands at 2 4 0 0 - 2 6 5 0 m o n the highest peak, the summit of K i m h a n d u . Luxuriance varies from a n almost impenetrable thicket of stems as thick as a finger and only 4 m tall o n shallow soil as at the foot of Sabinio, to well-spaced stems 8 c m in diameter and 15 m tall between which it is easy to walk, as o n the south-west slopes of M t Elgon. Flowering is gregarious, though rarely simultaneously over large areas. After flowering the whole plant dies and regeneration is by seed. Individual stems Uve for 5-10 years and the interval between flowering is believed to b e at least 30 years. It is possible that the trees which are often found scattered in b a m b o o become established at these times, w h e n the dead patches are covered with a strong growth of Rubus, Sambucus africana, Lobelia bambuseti and Impatiens. T h e extent to which Arundinaria has been able to occupy forest sites because of fire is uncertain. O n the M a u Range, according to Glover & T r u m p (1970), it is fire-induced, and charred stumps of Juniperus can frequently b e seen sticking through it. Elsewhere o n the M a u , Arundinaria forms an understorey in Juniperus forest and here too there is evidence of past burning. T h e most frequent trees which occur scattered in Arundinaria alpina b a m b o o are Afrocrania volkensii, Dombeya goetzenii, Faurea saligna, Hagenia abyssinica, Ilex mitis, Juniperus procera, Lepidotrichilia volkensii, Nuxia congesta, Podocarpus latifolius, Prunus africana, Rapanea melanophloeos and Tabernaemontanajohnstonii.
Afromontane evergreen bushland and thicket (mapping units 19a, 65, and 66) Refs.: Chapman & White (1970, p. 138, 148, 169-70); Edwards (1967, p. 189-90); Greenway (1955, p. 560; 1965; 1973, p. 55-6); Hedberg (1951); Killick (1963, p. 41-4, 80-4); Langdale-Brown et al. (1964, p. 33, 109); Lebrun (1942, p. 65-8); Lewalle (1972, p. 146-9); Lind & Morrison (1974, p. 145); Phipps & Goodier (1962, p. 306-7); Pócs (1974; 19766, p. 488-9); Richards (19636); White (1978a; M S , 1949, 1963, 1973, 1975-76). Phots.: Chapman (1962: 14); Chapman & White (1970: 52); Edwards (1967: 120); Hedberg (1951: lb); LangdaleBrown et al. (1964: 2); Lebrun (1942: 27b, 28a, 29a); Pócs (1974: 8). Profile: Lewalle (1972: 28). Syn.: Cave Sandstone scrub (Killick, 1963); Ericaceae-Stoeèe high montane heath (Langdale-Brown et al.); Ericaceous wooded grassland (Lind & Morrison, 1974); PasserinaPhilippia, Widdringtonia fynbos (Killick, 1963); Philippia forest (Hedberg, 1951); subalpine elfin forest (Pócs, 19766); upland moor p.p. (Greenway, 1973). Afromontane evergreen bushland and thicket occur o n most of the higher African mountains. They are also found o n the crests and summits of s o m e smaller mountains, especially those which are situated close to the sea or a large lake. Characteristically they occupy a large part of the 'Ericaceous belt' of Hedberg (1951). T h e y also occur locally o n shallow soils inside the m o n t a n e forest belt, and o n the exposed summits of
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Vegetation of the floristic regions
mountains too low to support an Ericaceous belt. They vary greatly infloristiccomposition but s o m e m e m b e r s of the Ericaceae (species of Blaeria, Erica, Philippia and Vaccinium) are nearly always present, and are sometimes exclusively dominant. Ericaceae are virtually absent, however, from the wetter types of elfin thicket. W h e r e the ground is not very rocky and has been protected from fire for several years the dominants form almost impenetrable thickets. These conditions are found on the wetter mountains such as Ruwenzori, where the tallest thickets occur. O n drier, rocky slopes the vegetation is often discontinuous and the bushes form an open canopy. Ericaceous bushland and thicket burn readily, and, especially o n the drier mountains, they have been extensively replaced by secondary grassland. Unburnt Ericaceous bushland and thicket is normally between 3 and 13 m tall. O n shallow soils and exposed slopes, however, they merge into Afromontane shrubland. Tall elfin thicket (3-7 m high) occurs o n the crests of certain low crystalline mountains in East African which rise steeply from the lowlands and are situated relatively close to the sea. They are too low to support an Ericaceous belt but their summits are in the zone of permanent mist. O n B o n d w a Peak (2120 m ) in the Uluguru M t s , Tanzania, the estimated m e a n annual rainfall is 3000 m m (Pócs, 1974). Here Syzygium cordatum is the most abundant of the larger w o o d y species. Its gnarled and semi-prostrate habit is totally different from the erect growth it shows lower d o w n the mountain. Afromontane and Afroalpine shrubland (mapping units 19a, 65, and 66) Refs.: Chapman (1962, p. 23); Chapman & White (1970, p. 170); Greenway (1955, p. 560-2); Jackson (1956, p. 370-1); Killick (1963, p. 78, 92-3); Phipps & Goodier (1962, p. 306 -8). Phots.: Chapman (1962: 23); Greenway (1955: 3); Killick (1963:27,34,44-6). Syn.: Erica-Helichrysum heath (Killick, 1963); open upland moorland (Greenway, 1955). O n shallow soils, and especially o n exposed rocky ridges, o n the high African mountains the Ericaceous bushland and thicket described above give w a y to m u c h shorter Afromontane shrubland, in which Ericaceae still usually play an important role. These short shrublands are very mixed communities in which, besides shrubs, grasses, Cyperaceae, forbs (especially geophytes), bryophytes and lichens are conspicuous. Afromontane shrubland consists partly of stunted individuals of the dominants of Ericaceous bushland and thicket, and partly of species which are normally absent from the latter c o m munities. Patches of dwarf shrubland also occur at higher altitudes o n the highest mountains as part of the Afroalpine mosaic. Afromontane and Afroalpine grassland (mapping units 19a, 65, and 66) Refs.: Chapman (1962); Chapman & White (1970); Fries & Fries (1948, p. 24-7); Greenway (1955, p. 555-8);
Hedberg (1964, p. 114-18); Herlocker & Dirschl (1972); G . Jackson (1969); J. D . Jackson (1956, p. 361-3, 368-9); Killick (1963); Lind & Morrison (1974, p. 150-1); Maitland (1932); Phipps & Goodier (1963); Pócs (1976ft, p. 494); Richards (19636, p. 548-53); V a n Zinderen Bakker & Werger (1974); White (1978a, p. 495-8, 504; M S , 1973, 1975-76); W o o d (1965). Phots.: Chapman & White (1970: 1-4, 9-10, 13-15, 19-20, 28-30, 32, 38-9); Greenway (1955: 4); Hedberg (1964: 10, 13, 14, 18, 19, 30, 69); Killick (1963: 2, 8, 21); Maitland (1932: 6, 7). Grassland, today, is the most widespread vegetation type o n the African mountains, especially the drier ones. There are undoubtedly s o m e small areas of edaphic grassland and it is equally certain that, in the absence of h u m a n interference, grassland could be maintained by naturally occurring fires, which result from lightning, landslides, or volcanic activity. T h e former extent of such edaphic and natural fire-climax grassland is a matter of controversy. There is n o w , however, n o reasonable doubt that most Afromontane grasslands have originated, or have been greatly extended, relatively recently as a result of m a n ' s destructive activity. O n the high mountains of tropical Africa the secondary grasslands of the Ericaceous and Afroalpine belts are quite different from those of the Forest belt in their floristic composition and chorological relationships. M o s t of the species which dominate secondary grassland in and above the Ericaceous belt belong to the tribes Festuceae, Aveneae, and Agrosteae, and are Afromontane (including Afroalpine) endemics, or, at least in Africa, are confined to the high mountains. They are normal constituents of Ericaceous and mixed Afroalpine communities or grow o n rocky slopes or in boggy hollows and are not dependent o n fire for their presence. Indeed several of them are intolerant of fire. B y contrast, nearly all the species which dominate secondary grassland in the Forest belt belong to the tribes Andropogoneae and Paniceae, and are also widespread in the African lowlands. They m a y have invaded the forest belt from the lowlands following the destruction of forest, or they m a y formerly have occurred within the forest zone as marginal intruders in a few places where forest w a s excluded b y unfavourable edaphic conditions. O n the tropical mountains this distinction between the two chorological elements is well marked. In South Africa, however, it is partly obscured because the 'temperate' genera descend m u c h lower and the most abundant 'tropical' species, Themeda triandra, ascends relatively higher. In general, secondary grassland develops m o r e readily o n the drier mountains, but other factors are sometimes important. Steep well-drained slopes such as those formed by the porous lava flows of M t C a m e r o u n , a wet mountain, are covered with secondary grassland. Readily combustible communities such as those d o m i nated by Ericaceae or conifers are m o r e vulnerable to fire than most types of broad-leaved forest, and have been m o r e extensively replaced than the latter.
The Afromontane archipelago-like regional centre of endemism and the Afroalpine archipelago-like region
Secondary montane grassland is sometimes invaded by small fire-resistant trees. Species of Protea are the most characteristic and occur o n African mountains from Ethiopia to South Africa. They are usually small bushy trees 3-5 m tall with very short twisted boles and thick deeply fissured bark. W h e r e fires are severe they are often fire-trimmed and there is n o regeneration. Persistent severe fire eliminates them completely. W h e n secondary montane grassland is protected from fire for several years it is eventually invaded by forest-precursor shrubs and climbers, which form a dense thicket. T h e latter suppresses the grass and is itself invaded b y secondary forest trees, which sometimes invade the protected grassland direct. M o s t examples of this succession are to be found in forest reserves where a deliberate policy of fire-protection has been followed. A similar development towards forest can, however, occur w h e n landslips provide fire-protected niches in otherwise annually burnt grasslands ( C h a p m a n & White, 1970, p . 139, phot. 34). T h e commonest grasses in secondary grassland in the forest belt o n tropical mountains are Elionurus argentens, Exotheca abyssinica, Loudetia simplex, Monocymbium ceresiiforme, Themeda triandra, and species of Andropogon, Brachiaria, Digitada, Hyparrhenia, Pennisetum, and Setaria. In the Ericaceous and Afroalpine belts they are largely replaced by species of Agrostis, Deschampsia, Festuca, Koeleria, Pentaschistis, and Poa. In the Natal Drakensberg above the forest belt the commonest species are Bromus speciosus, Festuca costata, Pentaschistis tysonii and Themeda triandra.
M i x e d Afroalpine communities (mapping units 65 and 66) In tropical Africa Refs.: H a u m a n (1933, 1955); Hedberg (1951-69; 1975); Mabberley(1973, 1974, 1976); Salt (1954). Phots.: Hedberg (1964: 6-109); Salt (1954: 6-9). Profiles: Hedberg (1964: 84, 96, 102, 104). T h e vegetation of the highest mountains of tropical Africa, those which reach altitudes between 3800 and 6000 m , namely Ruwenzori, the Virunga Voléanos, Elgon, Aberdare, M t K e n y a , Kilimanjaro and M t M e r u , is so different from that occurring at lower altitudes that it has attracted the attention of travellers and scientists since the earliest days of botanical exploration. It is characterized b y the occurrence of Giant Senecios (Senecio subgen. Dendroseneció), Giant Lobelias, shrubby Alchemillas and other plants of remarkable lifeform. In recent years the flora and vegetation of these
169
high peaks have been regarded by those with specialist knowledge ( H a u m a n , 1955; Hedberg, 1965) as being sufficiently distinct to justify the recognition of a separate phytogeographical Afroalpine Region, which Hedberg (1961) extended to include the higher peaks of Ethiopia. If the Afroalpine Region is delimited to coincide with the Afroalpine belt (Hedberg, 1951) then its totalflorais small (c. 280 species o n the East African mountains, though there are others in Ethiopia which are still insufficiently k n o w n ) . In the Afroalpine belt there are virtually n o endemic genera, and very few species which d o not also occur in the Ericaceous and forest belts. For these reasons White (1978a) has suggested that the Afroalpine and Afromontane Regions should be combined, though for certain purposes the former can be recognized as an archipelago-like region of extreme floristic impoverishment. Afroalpine vegetation is physiognomically very mixed and does not readily fit the major physiognomic categories. Hedberg, o n whose scholarly studies our knowledge of Afroalpine vegetation is almost entirely based, recognizes five distinctive Afroalpine life-forms, all of which are closely paralleled in other genera in the ' p a r a m o ' vegetation of the northern Andes in South America. These five life-forms (and other less-specialized types) combine kaleidoscopically to give an infinite variety of mixtures. Vegetation consisting exclusively of a single life-form occupies only small areas or extreme habitats. T h e tropical African islands of Afroalpine vegetation are too small to show o n the m a p . They are included with the Ericaceous belt as part of mapping unit 65 (Altimontane vegetation). In South Africa Refs.: Coetzee (1967); Killick (1978a, 19786, 1978c); Van Zinderen Bakker & Werger (1974). Phots.: Killick (1978c: 14-18); Van Zinderen Bakker & Werger (1974: 2-4). T h e authors cited above place in the Afroalpine Region all vegetation above the forest belt in the southern Drakensberg. This is at variance with Hedberg's treatm e n t for tropical Africa since he excludes the Ericaceous belt. Although there are s o m e resemblances between East and South Africa there are also important differences, notably the absence of giant Lobelias and Senecios from the latter. Killick has reviewed the Afroalpine Region in southern Africa (1978c) and described its vegetation for one part in great detail (1963). In the present work the vegetation of the crest of the Drakensberg is referred to as 'Altimontane' (mapping unit 66).
x
The Guinea-Congolia/Zambezia regional transition zone
Geographical position and area
Geographical position and area
Geology and physiography
The transition zone which separates the GuineoCongolian and Zambezian Regions extends from the Atlantic Ocean to the high ground flanking the northern end of Lake Tanganyika. Its m a x i m u m width is nearly 500 k m . (Area: 705000 k m 2 . )
Climate Flora Mapping units Vegetation Drier peripheral semi-evergreen Guineo-Congolian rain forest Zambezian dry evergreen forest and transition woodland Grassland and wooded grassland The coastal mosaic
Geology and physiography Most of the transition zone forms part of the dissected plateau which extends south from the Zaire basin to the Zambezi-Zaire watershed. T h e western limit of the plateau is marked by a well-defined escarpment cut into Precambrian rocks. There is a narrow coastal plain of Cretaceous and more recent sediments which is separated from the escarpment by a somewhat wider belt of undulating but low-lying country underlain by Precambrian rocks. M u c h of the plateau in eastern Angola and K w a n g o is overlain by a thick mantle of Kalahari Sand, but in the deeply entrenched river valleys which mostly run in a north-south direction the underlying Karoo strata are exposed. Further east in Kasai, Karoo beds predominate in the north and Precambrian rocks in the south, but the long narrow ridges between the valleys are mostly covered with Kalahari Sand. Further east still, Precambrian rocks predominate. The altitude of the plateau is mostly between 1000 and 1500 m .
Climate In most places the climate is intermediate between those of the Guineo-Congolian and Zambezian Regions. T h e dry season is more severe than that of the former but less so than that of the latter. Rainfall diminishes very rapidly near the Atlantic coast to below 800 m m per year, but dry-season relative humidity is high. Frost is u n k n o w n in the transition zone. (See Fig. 15.)
Flora Excluding marginal intruders there are probably no more than 2000 species, very few of which are endemic.
The Guinea-Congolia/ Zambezia regional transition zone
171
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172
Vegetation of the floristic regions
O f the 841 species recorded from the Kaniama District transition zone climate changes rapidly from the arid of Lower Shaba by Mullenders (1954, 1955), 31.3 per coastal plain, where the vegetation is predominantly cent are Guineo-Congolian species and 31.4 per cent are Zambezian, to the humid, essentially GuineoZambezian ('Sudano-Zambezian') species. Most of the Congolian, D e m b o s 'cloud' forests on the escarpment of remainder are linking species with wider distributions. the interior plateau. Only 14 species (1.6 per cent) are endemic. In addition to the vegetation described below, Other endemic species which are confined to the m i o m b o woodland, similar to that described in Chaptransition zone or only slightly transgress its boundaries ter II, occurs in the southern part, but it is floristically include Combretum camporum, Crotón dybowskii, poor and little is k n o w n about it. Diospyros grex, D. heterotricha, D. wagemansii, Hymenostegia laxiflora, Pteleopsis díptera and Rinorea malemDrier peripheral semi-evergreen baensis. Most of these only occur near the coast. Guineo-Congolian rain forest (mapping units 2 , 11a, and 14)
Mapping units 2 (p.p.). Drier peripheral semi-evergreen GuineoCongolian rain forest. 11a (p.p.). Mosaic of Guineo-Congolian rain forest and secondary grassland. 14. Mosaic of Guineo-Congolian rain forest, Zambezian dry evergreen forest and secondary grassland. 15 (p.p.). West African coastal mosaic. 21 (p.p.). Mosaic of Zambezian dry evergreen forest, wetter m i o m b o woodland and secondary grassland. 31. Mosaic of wetter Zambezian woodland and secondary grassland (see page 61). 37 (p.p.). Acacia polyacantha secondary wooded grassland (see page 63). 60 (p.p.). Edaphic and secondary grassland on Kalahari Sand.
Vegetation Réf.: White & Werger (1978). The floras of the Guineo-Congolian and Zambezian Regions are almost mutually exclusive. There is, however, a transition zone between them up to 500 k m wide and of considerable complexity. In it an impoverished Guineo-Congolian flora and an even more impoverished Zambezian flora interdigitate or occur in mosaic, and locally intermingle. T h e greater part of the transition zone is occupied today by secondary grassland and wooded grassland dominated almost exclusively by Zambezian species. The latter, however, have greatly increased in abundance following destruction of the original vegetation, but it seems that formerly they occurred in most parts of the transition zone, though often confined to edaphically specialized sites such as rocky places and seasonally waterlogged grassland. In general, the Zambezian element becomes more abundant towards the south, and, where soil permits, the transition forms a continuum. Elsewhere, edaphic conditions override the climate to produce a mosaic. A t the western end of the
Forest of Guineo-Congolian type has an uneven distribution in the transition zone. T h e most extensive of the surviving forests are the D e m b o s 'cloud' forests in Angola. They occur between 350 and 1000 m about 100 k m from the sea on the escarpment leading up to the plateau. Rainfall is between 1100 and 1500 m m per year, but the existence of forest is largely dependent on the constant condensation of water-vapour brought in by moisture-laden sea breezes. The surrounding country is too dry for the occurrence of forest. Further west in north-west Angola and Zaire vast tongues of Guineo-Congolian forest penetrate towards the south in the wide valleys of the major tributaries of the Zaire River. These forests are not confined to the banks ofriversbut also occur on the fertile soils of the rejuvenated land surface and are often several kilometres wide. Further west still, in the K a n i a m a region of Lower Shaba, rain forest of Guineo-Congolian affinity was formerly the prevalent vegetation on soils derived from gabbro and tonalité. All that survives today are narrow bands on the slopes of small valleys which have cut into the plateau, and a few isolated remnants on the plateau itself (Mullenders, 1954). Larger tree species, not all of which are generally distributed, include Albizia zygia, Antiaris toxicaría, Trilepisium madagascariense, Canarium schweinfurthii, Celtis zenkeri, Chlorophora excelsa, Cynometra alexandri, Dacryodes edulis, Entandrophragma angolense, Khaya anthotheca, Klainedoxa gabonensis, Lovoa trichilioides, Pachystela brevipes, Parkia filicoidea, Petersianthus macrocarpus, Piptadeniastrum africanum, Pycnanthus angolensis, Ricinodendron heudelotii, Staudtia stipitata and Treculia africana. Zambezian dry evergreen forest and transition woodland (mapping units 14 and 21) Dry evergreen forest of pronounced Zambezian affinity is widely distributed on the Kalahari Sand-covered plateau of K w a n g o , where it is k n o w n as 'mabwati', and in adjacent parts of Angola. It is up to 25 m tall. T h e leaves of the trees are more coriaceous than those of
The Guinea-Congolial Zambezia regional transition zone
rain-forest species and lack 'drip tips'. T h e most characteristic taller trees in the mabwati described b y Duvigneaud (1950, 1952) are Marquesia macroura, M . acuminata, Berlinia giorgii, Latinea antiscorbutica, Daniellia alsteeniana, Brachystegia spiciformis, B. wangermeeana and Parinari curatellifolia. Smaller trees include Uapaca nítida, U. sansibarica, Memecylon sapinii, Diospyros batocana, Anisophyllea gossweileri, Monotes dasyanthus and Diplorhynchus condylocarpon. The two species of Marquesia, Berlinia giorgii and Daniellia alsteeniana, only occur in the Guinea-Congolia/Zambezia transition zone and in the wetter northern parts of the Zambezian Region. Most of the other species are very widely distributed in the Zambezian Region. T h e floristic composition Usted above suggests transition woodland (page 91) or a degraded community rather than true forest. T h efloristiccomposition of mabwati changes from north to south. In the north of K w a n g o , in the KalahariKaroo contact zone, there is a strong admixture of Guineo-Congolian species, especially those which also occur o n Kalahari Sand in the teke forests of the Brazzaville-Kinshasa region. S o m e of the dominants of mabwati, including Daniellia alsteeniana, Marquesia acuminata and M . macroura, are also widely distributed to the west of K w a n g o in the Malanje Province of Angola (mapping unit 14), though there is little published information. T h e m i o m b o dominants seem to be virtually absent from this region. Tall transition woodland (8-10 m ) , dominated by Berlinia giorgii and Uapaca nitida and related to the mabwati of K w a n g o , occurs in the K a n i a m a region o n granite outcrops where it forms a zone between forest of Guineo-Congolian affinity in the valleys, and a sparsely wooded grassland of Zambezian affinity o n the plateau above (Mullenders, 1954). Berlinia giorgii is the most abundant species, followed b y Uapaca nitida and Combretum psidioides. Other Zambezian species such as Albizia versicolor, Cussonia sessilis, Maprounea africana, Monotes dasyanthus, Piliostigma thonningii, Sterculia quinqueloba, Stereospermum kunthianum, Strychnos cocculoides and Terminalia mollis are rarer. Although fire rarely passes through the Berlinia giorgii forests they show little tendency to change in the direction of Guineo-Congolian forest. Guineo-Congolian linking species are few, e.g. Canarium schweinfurthii, and mostly occur o n termite m o u n d s . Grassland and wooded grassland (mapping units 11a, 14, 34, 37, and 60) These communities are mostly secondary but some small patches m a y be primary. They vary greatly in floristic composition and luxuriance, chiefly in relation to parent material, catenary position, and degree of degradation. S o m e general information is given o n page 50. T h e grasslands occurring in Lower Zaire have been described in considerable detail by Duvigneaud (1952). Those of
173
K w a n g o and the Kaniama region of Shaba are briefly considered below. In K w a n g o the mabwati forests of the Kalahari Sand-covered plateau have largely been replaced by a wooded grassland ('mikwati') in which the most frequent fire-resistant trees are Erythrophleum africanum, Dialium engleranum, Burkea africana, Hymenocardia acida, Diplorhynchus condylocarpon, Pterocarpus angolensis, Protea petiolaris, Combretum celastroides subsp. laxiflorum and Strychnos pungens. T h e herb layer consists largely of the grasses Hyparrhenia diplandra, H. familiaris, Loudetia arundinacea, Digitaria diagonalis (uniglumis), Brachiaria brizantha and Ctenium newtonii. O n the deepest sands, which m a y be more than 100 m thick, the structureless soil is without h u m u s and is extremely deficient in nutrients. Although the rainfall is 1600-1800 m m per year, it percolates rapidly and the water-table is at a considerable depth. Fires occur each year and have been responsible for degrading the vegetation to a sparse short grassland which includes m a n y geophytes and geoxylic suffrutices, and is referred to by Belgian authors as 'steppe', 'pseudosteppe' or 'savane steppique'. According to Devred et al. (1958) these grasslands are secondary and have relatively recently replaced forest and woodland. It is likely, however, that at least locally where the water-table is high for part of the year, a similar community represents an edaphic climax from which the species of secondary grassland have been recruited. In K w a n g o the principal grasses of secondary suffrutex grassland are Aristida vanderystii, Ctenium newtonii, Digitaria brazzae, Diheteropogon grandiflorus (emarginatus), Elionurus argenteus, Loudetia demeusii, L simplex, Monocymbium ceresiiforme, Rhynchelytrum amethystinum, Schizachyrium thollonii and Tristachya nodiglumis (eylesii). T h e geoxylic suffrutices include Anisophyllea quangensis, Brackenridgea arenaria, Erythrina baumii, Gnidia kraussiana, Landolphia camptoloba, Ochna manikensis, Parinari capensis and Rauvolfia nana. These grasslands are sometimes lightly wooded with scattered trees of Combretum, Dialium engleranum, Erythrophleum africanum, and Daniellia alsteeniana, but often are treeless for considerable distances, or contain only stunted shrubby individuals of Swartzia madagascariensis, Burkea africana, Oldfieldia dactylophylla and Hymenocardia acida. T h e secondary grasslands of the K a n i a m a region of Shaba have been described b y Mullenders (1954). Similar grassland extends for 300 k m to the north. T h e flora of the secondary grasslands near K a n i a m a is extremely poor, consisting of a mere 252 species, of which 19 are grasses. A few forest pioneers, such as Albizia adianthifolia, Clausena anisata, Harungana madagascariensis and Phyllanthus muelleranus, are Guineo-Congolian linking species but the remainder, almost without exception, are Zambezian species some of which also occur in other savanna regions. T h e most notable grasses are Andropogon schirensis (dominant o n the more degraded sites), Hyparrhenia
174
Vegetation of the floristic regions
are also typical members of secondary wooded confinis (dominant o n the less-degraded sites), Pennigrassland. setum unisetwn, Brachiaria brizantha, Digitaria diagonalis, Elymandra androphila, Hyparrhenia filipéndula, H . newtonii (lecomtei), H . rufa, Hyperthelia dissoluta {HyparT h e coastal mosaic rhenia ruprechtii), Imperata cylindrica, Loudetia arundi(mapping unit 15) nacea, Panicum baumannii (fulgens), P. phragmitoides, Schizachyrium brevifolium and Urelytrum giganteum. T h e rainfall of the coastal belt to the west of the cloud forests of D e m b o s and northwards into Lower Zaire and T h e Zambezian w o o d y flora is represented by the Cabinda is too low to support Guineo-Congolian following heliophilous, pyrophytic trees: Acacia hockii, vegetation except along watercourses. A. polyacantha subsp. campylacantha, A. sieberana, Albizia versicolor, Annona senegalensis, Bridelia ferruT h e prevalent vegetation is grassland and wooded ginea, Dombeya shupangae, Erythrina abyssinica, Gargrassland, most of it probably secondary. Adansonia denia temifolia, Grewia mollis, Hymenocardia acida, digitata, which is rare or absent further inland, is a Maprounea africana, Maytenus senegalensis, Monotes conspicuous feature of the landscape. T h e two caloneurus, M . mutetetwa, Ochna schweinfurthiana, Pariintroduced trees, Anacardium occidentale and Mangifera nari curatellifolia, Pericopsis angolensis, Piliostigma thonindica, are also plentiful. O n sandy soil, o n elevated ningii, Psorospermum febrifugum, Pterocarpus angolensis, country of the littoral, where there is regular Schrebera trichoclada, Sclerocarya caffra, Secwidaca condensation of moisture from the westerly winds, longepedunculata, Sterculia quinqueloba and StereoStrychnos henningsii forms dense thickets thousands of spermum kunthianum. square kilometres in extent. Granite outcrops near Matadi support a special vegetation rich in lichens and There is s o m e evidence that m a n y of the Zambezian succulents such as Sansevieria cylindrica, Aloe, Rhipsalis species which today dominate secondary wooded grassand Euphorbia. land in the K a n i a m a region have been recruited from edaphically specialized refugia in which the Zambezian A t the m o u t h of the Zaire River there are 250 k m 2 of flora has probably persisted from a drier epoch. Thus, s w a m p forest o n the landward side of the mangrove. Mullenders describes a Loudetia arundinacea, Ochna Although the annual rainfall is n o more than 700 m m , leptoclada wooded grassland from the upper slopes and the constituent species are almost exclusively Guineoridges of granite outcrops which appears to be an Congolian rain-forest species and some, e.g. Sacoglottis edaphic climax. M a n y of the trees, including gabonensis, are especially characteristic of the wetter Stereospermum kunthianum, Entada abyssinica, Parinari types. Elaeis guineensis, besides occurring in this c o m curatellifolia, Sterculia quinqueloba, Albizia versicolor,munity, forms natural societies further south in very hot Sclerocarya caffra and Terminalia mollis, which occur sheltered situations where it is dependent o n phreatic a m o n g piles of rocks or rooted in the granite pavement, water, since annual rainfall is only 600 m m .
xi
The Guinea-Congolia/Sudania regional transition zone
Geographical position and area
Geographical position and area
Geology and physiography
This transition zone, which separates the G u i n e o Congolian and the Sudanian Regions, extends across Africa from Senegal to western U g a n d a . Between eastern G h a n a and Benin Republic ( D a h o m e y ) it reaches the coast, where the well-known ' D a h o m e y gap' separates the Guineo-Congolian rain forests into two blocks of unequal size. T h e vegetation of the driest parts of the coastal strip is not truly transitional but is included here for convenience. (Area: 1165000 k m 2 . )
Climate Flora Mapping units Vegetation The Coastal Plain of Ghana West African dry coastal forest Seasonally waterlogged grassland on the Accra Plains Termite-mound thicket The Coastal Plain of Basse Casamance
Geology and physiography Nearly everywhere the altitude is less than 750 m . In places the C a m e r o u n Highlands rise to more than 2000 m but their highest peaks belong to the Afromontane archipelago. Otherwise the land reaches more than 1000 m only in Fouta Djalon, the Guinea Highlands and the Togo-Atacora range, which all extend into the Guineo-Congolian Region, and on the Jos Plateau in Nigeria, which is shared with the Sudanian Region. T h e Jos Plateau is the largest area of land in Nigeria over 1200 m altitude. F r o m its surface some rocky hills rise 150 to 300 m above this. It is composed principally of granite and basalt. T h e former is chiefly responsible for its rugged topography, including the steep escarpment up to 600 m high on the west and south. T h e geology of this transition zone is very diverse. Palaeozoic rocks predominate in Guinea-Bissau and the Volta basin. In Nigeria there are extensive areas of Cretaceous sediments, especially in the Benue and lower Niger valleys. Most other places are underlain by the Precambrian.
Climate Nearly everywhere the climate is transitional between those of the Guineo-Congolian and Sudanian Regions. A narrow strip of coastal plain in West Africa extending from G h a n a eastwards to Benin Republic has, however, an anomalously dry climate. In the driest part, near Accra, rainfall is only 733 m m per year. T h e dryness of the plains is further accentuated by the desiccating action of strong onshore breezes which blow throughout
176
Vegetation of the floristic regions
the year. Because of w i n d action forest is restricted to the protected leeward slopes of hills, thicket c l u m p s are elongated in the direction of the prevailing winds a n d the c r o w n s of isolated trees are severely pruned (Jenik & Hall, 1976). The controlling effect of the harmattan wind on vegetation in the Togo Mts in Ghana, which he astride the northern b o u n d a r y of the transition zone, has been described b y Jenik a n d Hall (1966). (See Fig. 16.)
Flora There are probably fewer than 2000 species, nearly all of which are Guineo-Congolian or Sudanian wides, or linking species with even wider distributions. T h e upland areas of Guinea Republic and adjacent Sierra Leone between 700 and 1000 m , however, support a few endemic species including Bafodeya benna and Fleurydora felicis, which both belong to monotypic genera, and Diospyros feliciana. A few Afromontane species also occur in Fouta Djalon in Guinea Republic. The Accra Plains, for their size, harbour a remarkable concentration of endemic and disjunct species (Jenik & Hall, 1976). The former include Commiphora dalzielii, Grewia megalocarpa, Talbotiella gentii a n d Turraea ghanensis. A m o n g the disjuncts, Crossandra nilotica a n d Ochna ovata elsewhere occur only in East Africa. Other disjuncts, including Capparis fascicularis, Grewia villosa a n d the grasses Aristida sieberana, Chloris prieurii a n d Schoenefeldia gracilis, have their m a i n areas in the Sahel a n d northern S u d a n zones far to the north.
Mapping units 2 (p.p.). Drier peripheral semi-evergreen G u i n e o Congolian rain forest (see Chapter I). 11a (p.p.). M o s a i c of Guineo-Congolian rain forest a n d secondary grassland (see Chapter I). 12. M o s a i c of Guineo-Congolian rain forest, I sober linia w o o d l a n d a n d secondary grassland. 13. M o s a i c of G u i n e o - C o n g o l i a n rain forest, secondary grassland a n d m o n t a n e elements. 15 (p.p.). W e s t African coastal mosaic.
Vegetation Refs.: Clayton (1961); Keay (1948, 1959a, 1959c). T o d a y the greater part of the Guinea-Congolia/Sudania transition zone is covered with secondary grassland a n d secondary w o o d e d grassland similar to that described in Chapter I. Various types of forest were formerly widespread, but they have b e e n extensively destroyed b y fire a n d cultivation. O f the surviving remnants the m o s t luxuriant are indistinguishable from the drier types of peripheral semi-evergreen rain forest (page 79), a n d
indeed represent a northern extension or outliers of it. In addition, shorter, floristically poorer forest also occurs. A t o n e time it w a s thought that virtually the whole of the transition zone h a d been covered with forest. It n o w seems likely, however, that patches of Isoberlinia w o o d land a n d Monotes w o o d l a n d similar to s o m e of the m o s t characteristic communities of the Southern S u d a n zone (page 106) formerly occurred o n shallow soils, a n d that transition w o o d l a n d (page 105) formed the ecotone between forest a n d w o o d l a n d . M o s t of the forest formerly occurring in the uplands of F o u t a Djalon (mapping unit 13, alt. c. 1 0 0 0 - 1 5 0 0 m ) has b e e n replaced b y cultivation a n d secondary grassland. A s in other parts of the U p p e r G u i n e a highlands, the m o s t abundant tree in the forest is Parinari excelsa (page 81). There are also s o m e A f r o m o n t a n e species such as Nuxia congesta, but they are less n u m e r o u s than in the m o r e elevated parts of the highlands further east. T h e m o s t characteristic species of s w a m p forest and riparian forest in the western half of the G u i n e a Congolia/Sudania transition zone are Berlinia grandiflora, Cola laurifolia, Cynometra vogelii, Diospyros elliotii, Parinari congensis, a n d Pterocarpus santalinoides. T h e y all also penetrate s o m e distance into the G u i n e o - C o n g o l i a n Region or are extensively distributed within it. T h e Guinea-Congolia/Sudania transition zone includes both the Derived S a v a n n a zone a n d the Southern G u i n e a zone of K e a y (1959a). It corresponds quite closely to the 'zone des savanes subforestières avec galeries' of Chevalier (1938) w h i c h separates his 'zone soudanaise proprement dite' from his 'zone nord de la grande forêt dense'. Within the transition zone there is n o simple relationship between latitude a n d the extent to w h i c h the original forest has survived. This is partly because rainfall does not always s h o w a regular diminution northwards. Also the original vegetation w a s profoundly influenced b y parent material, a n d its m o d i fication b y m a n has been closely related to population density. T h e coastal plain of G h a n a a n d that of Basse C a s a m a n c e have distinctive local features a n d are described separately below. T h e Coastal Plain of G h a n a ( m a p p i n g unit 15) In the driest part of the plain, near Accra, the m o s t extensive soils are unfavourable to tree growth a n d support a sparse short grassland, only 8 0 c m tall, dotted with thicket c l u m p s which occur o n low flat m o u n d s . Elsewhere various types of forest, less luxuriant than G u i n e o - C o n g o l i a n rain forest, represent the climax. West African dry coastal forest Refs.: Hall & Swaine (1974; 1976); Jenik & Hall (1976, p. 203-4). Phot.: Jenik & Hall (1976: 3). Profiles: Hall & Swaine (1976: 15); Jenik & Hall (1976: 8).
The Guinea-Congolia/Sudania
regional transition zone
%&5B
177
Vegetation of the floristic regions
178
T h e r e are t w o types of evergreen or semi-evergreen forest. In the western type (J. B . Hall, pers. c o m m . ) there are t w o or three tree layers. T h e m a i n canopy is at 15-20 m a n d is characterized b y Cynometra megalophylla and Manilkara obovata, which are m u c h m o r e abundant in this type than in rain forest. Emergents u p to 3 0 m tall of Nesogordonia papaverifera, Celtis mildbraedii, Antiaris toxicaría a n d Ceiba pentandra often occur. Lianes are abundant, b u t epiphytes are virtually absent a n d the field layer is poorly developed. T h e eastern type, which occurs o n inselbergs o n the A c c r a Plains, is always less than 2 0 m tall. Diospyros abyssinica and Millettia thonningii dominate the canopy, while the lower storey is c o m p o s e d chiefly of Drypetes parvifolia, D . floribunda a n d Vepris heterophylla. M o s t species are capable of coppicing a n d the undergrowth is extremely sparse. Large lianes are present, especially Griffonia simplicifolia a n d Premna cuadrifolia, but epiphytes other than lichens are lacking. Grasses are absent from the herb layer but a few succulents, e.g. Sansevieria liberica, occur. Seasonally waterlogged grassland on the Accra Plains Refs.: Jenik & Hall (1976). Phots.: Jenik & Hall (1976: 4, 5). T h e soils are shallow a n d parent rock is encountered within t w o metres of the surface. Drainage is i m p e d e d and root development, particularly of w o o d y plants, is inhibited. There are t w o m a i n types of soil, developed over acidic a n d basic gneiss respectively. T h e m o s t extensive soils overlying acidic gneiss are Pallid Sands with a n underlying impervious stone/clay p a n , which locally reaches the surface. T h e soils occurring over basic gneiss are Black Earths, which have a b u n d a n t calcium carbonate concretions a n d develop deep cracks in the dry season. T h e general d o m i n a n t of the drier grasslands of the Accra plains, both o n acidic a n d basic soils, is Vetiveria fulvibarbis. Its m o s t widespread associates are Brachiaria falcifera, Andropogon canaliculatus, Cassia mimosoides, Fimbristylis pilosa and Polygala arenaria. Termite-mound
thicket
Refs.: Jenik & Hall (1976); Okali et al. (1973). T h e m o u n d s are u p to 15 m in diameter but less than 0.5 m high. Termite activity is greater o n the m o u n d s than in the surrounding grasslands. Within the thicket patches there are small termitaria occupied b y Odontotermes pauperans a n d Amitermes evuncifer, a n d larger termitaria, u p to 3 m tall a n d 3 m wide at the base, occupied b y Macrotermes bellicosus. Although it is likely that the m o u n d s have b e e n built u p b y termite activity
over a long period of time, there is n o evidence that n e w m o u n d s are being formed in the grasslands today. O n the contrary, Okali et al. suggest that these clumps m a y be remnants of formerly continuous thicket. T h e patches of thicket have a dense closed canopy at 5 m , c o m p o s e d principally of Flacourtia indica (flavescens), Zanthoxylum zanthoxyhides, Grewia carpinifolia, Securinega virosa, Capparis erythrocarpos a n d Uvaria chamae. M a t u r e emergent trees of Elaeophorbia drupifera and Diospyros mespiliformis reach a height of 10 m . T h e succulent-leaved herb, Sansevieria liberica, is conspicuous in the field layer. T h e Coastal Plain of Basse C a s a m a n c e ( m a p p i n g unit 1 la, p.p.) Refs.: A d a m (1961a); Aubréville (19486). Forest of Guineo-Congolian affinity extends along the W e s t Coast of Africa into Basse C a s a m a n c e a n d Sénégal far b e y o n d the climatic limits of rain forest. In Basse C a s a m a n c e rainfall is 1 5 0 0 - 1 8 0 0 m m per year but m o s t falls in only five m o n t h s of the year a n d the dry season is too long a n d too severe to permit the development of typical rain forest. T h e country, however, is m o r e or less flooded during the rainy season a n d the water-table is not far beneath the surface during the dry season. This enables s o m e G u i n e o - C o n g o l i a n forest species to extend their range into a n inhospitable climate. M o s t of the natural vegetation has been replaced b yrice-fieldsor ground-nuts (Arachis), but until recently sufficient relicts remained to enable a reconstruction of the original vegetation to b e m a d e . Parinari excelsa w a s formerly a b u n d a n t both in permanent s w a m p forest in depressions a n d in drier forest o n better-drained soils. Forest o n the latter, which is about 18-20 m tall, is dominated b y Parinari excelsa. Other a b u n d a n t species are Erythrophleum suaveolens, Detarium senegalense, Afzelia africana and Khaya senegalensis. Rarer associates include Albizia adianthifolia, A . ferruginea, A . zygia, Antiaris toxicaría, Chlorophora regia, Cola cordifolia, Daniellia ogea, Dialium guiñéense, Monis mesozygia, Schrebera arbórea and Sterculia tragacantha. M o s t species of the drier Parinari-ErythrophleumDetarium forests of Basse C a s a m a n c e are also widely distributed in G u i n e o - C o n g o l i a n rain forest, especially in the drier semi-evergreen types. Although the C a s a m a n c e forest is without e n d e m i c species, it is a very distinct type, both floristicalry a n d in structure (Aubréville, 1949ft, p . 41). T h e c a n o p y is at 18-20 m a n d is m a d e u p of large trees which usually branch close to the ground a n d have leaning boles a n d very wide crowns. Trees with straight boles are rare, chiefly Khaya. Lianes of all sizes are abundant. T h e c a n o p y replaces its leaves before the e n d of the dry season, during which the 3-5 m tall understorey is mostly deciduous. T h o s e leaves which are not shed have a wilted appearance.
xii
T h e Lake Victoria regional mosaic
Geographical position and area
Geographical position and area
Geology and physiography
This Region includes most of U g a n d a , the whole of eastern R w a n d a and Burundi, and small parts of Zaire, K e n y a , and Tanzania. A small exclave occupies the Ruzizi Valley north of Lake Tanganyika. (Area: 224000 k m 2 . )
Climate Flora Mapping units Vegetation Drier peripheral semi-evergreen Guineo-Congolian rain forest Transitional rain forest Swamp forest Scrub forest Evergreen and semi-evergreen bushland and thicket and derived communities
Geology and physiography T h e Lake Victoria basin was formed during the middle Pleistocene b y earth movements associated with the evolution of the western a r m of the Great Rift Valley. F r o m the lake itself (altitude 1134 m ) the land surface falls gradually to the north. It rises gently to the south but m u c h m o r e abruptly to the east and west towards important islands of the Afromontane Region. Nearly everywhere the underlying rocks belong to the Precambrian but locally they are covered b y recent alluvium.
Climate Climatic gradients are often steep and are related to the complex physiography and to distance from Lake Victoria, which is an important source of precipitation. Locally the rainfall is sufficiently high (15002000 m m per year) and well distributed throughout the year to support rain forest. Elsewhere it is too low for rain forest but is not sufficiently seasonal for woodland. H e n c e scrub forest and semi-evergreen bushland and thicket represent the climax. A few m i o m b o species reach their northern limits near the southern shore of Lake Victoria. T h e dry season there is m u c h less severe than in most of the Zambezian Region and the communities in which m i o m b o species occur are not typically Zambezian. (See Fig. 17.)
Flora There are possibly n o m o r e than 3000 species, of which very few are endemic. There are probably n o endemic genera.
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Vegetation of the floristic regions
SERERE
(1139m) 24 1 1366
(5-25).
ENTEBBE (1182 m) 217 (15-50).
1505
500
F I G . 17. Climate and topography of the Lake Victoria regional mosaic (XII)
1000 Km
The Lake Victoria regional mosaic
Mapping units 2 (p.p.)- Drier peripheral semi-evergreen G u i n e o Congolian rain forest. 4 (p.p.). Transitional rain forest. 8 (p.p.). S w a m p forest. 11a (p.p.). Mosaic of Guineo-Congolian rain forest and secondary grassland. 25 (p.p.). Wetter Zambezian m i o m b o woodland (see Chapter II). 26 (p.p.). Drier Zambezian m i o m b o woodland (see Chapter II). 42 (p.p.). Somali-Masai Acacia-Commiphora deciduous bushland (see Chapter IV). 45 (p.p.). Mosaic of East African evergreen bushland and secondary Acacia w o o d e d grassland. Units 25, 26, and 42 reach their limits at the southern end of Lake Victoria. Their vegetation there is not very typical and little is k n o w n about it. It is not considered further here. M o s t of the species of secondary grassland and w o o d e d grassland in unit 1 la also occur in G u i n e o Congolian secondary grassland (page 84).
Vegetation T h e L a k e Victoria Regional Mosaic is the meeting-place of five distinct floras: Guineo-Congolian, Sudanian, Zambezian, Somalia-Masai and Afromontane. Its vegetation consists of a mosaic of floristically impoverished variants of the characteristic vegetation of the first four, in s o m e cases with an admixture of Afromontane species. Drier peripheral semi-evergreen Guineo-Congolian rain forest (mapping units 2 and 11a) Refs.: Eggeling (1947); Langdale-Brown, Osmaston & Wilson (1964, p. 44-51); G . H . S. W o o d (1960, p. 26-31). Phots.: Eggeling (1947: 1-4, 6, 7). Profiles: Eggeling (1947: 6-9); Langdale-Brown et al. (1964: 10). T h e majority of species are widespread in the G u i n e o Congolian Region. Large trees in this category include Albizia spp., Alstonia boonei, Aningeria altissima, Antiaris toxicaría, Chrysophyllum albidum, Celtis spp., Chlorophora excelsa, Cynometra alexandri, Entandrophragma angolense, E. cylindricum, E. utile, Holoptelea grandis, Khaya anthotheca, K. grandifoliola, Maesopsis eminii, Mildbraediodendron excelsum, Monts mesozygia (láctea), Piptadeniastrum africanum and Pycnanthus angolensis. Transitional rain forest (mapping unit 4) Refs.: Faden (1970); Lewalle (1972); Lucas (1968). Little has been published and only small fragments remain.
181
In western Burundi between 1600 and 1900 m the larger trees include Alangium chínense, Albizia gummifera, Anthonotha pynaertii, Carapa grandiflora, Chrysophyllum gorungosanum, Diospyros gabunensis, Newtonia buchananii, Parinari excelsa, Prunus africana, Strombosia scheffleri, Symphonia globulifera, Syzygium guiñéense and Xymalos monospora. These species also occur in western R w a n d a and eastern Kivu at the same altitudes (P. Bamps, pers. c o m m . ) . Kakamega forest in Kenya, which has fewer Afromontane elements, is shown on the m a p as lowland forest. Several Guineo-Congolian lowland rain-forest species, including Aningeria altissima, Cordia millenii, Entandrophragma angolense, Maesopsis eminii and Monodora myristica, reach their easternmost limit here. This forest, which is situated at 1520 to 1680 m o n the Nandi escarpment east of Lake Victoria, contains the following Afromontane species: Apodytes dimidiata, Macaranga kilimandscharica, Neoboutonia macrocalyx, Prunus africana, Strombosia scheffleri and Turraea holstii. S w a m p forest (mapping unit 8) Refs.: Eggeling (1935, p. 431; 1947, p. 60-1, 86-7); LangdaleBrown et al. (1964, p. 74-5); Lind & Morrison (1974, p. 5 0 8); G . H . S. W o o d (1960, p. 30-1). Phot.: Langdale-Brown et al. (1964: 24). Profile: Lind & Morrison (1974: 1.8). S w a m p forest dominated by species which are widespread in tropical Africa occurs extensively o n the shores of L a k e Victoria and elsewhere. T h e m o r e important species are Anthocleista schweinfurthii, Erythrina excelsa, Ficus congensis, Macaranga monandra, M . pynaertii, M . schweinfurthii, Mitragyna stipulosa, Musanga cecropioides, Parkia filicoidea, Pseudospondias microcarpa, Spondianthus preussii, Syzygium cordatum, Uapaca guineensis, Voacanga thouarsii and the palms Phoenix reclinata and Raphia farinifera. Acacia kirkii subsp. mildbraedii occurs sparingly in places. T h e forests occurring at c. 1200 m o n alluvial deposits at the m o u t h of the Kagera River o n the western shore of Lake Victoria are unique in tropical Africa in being composed in almost equal proportions of lowland, predominantly Guineo-Congolian, and Afromontane species. Undisturbed forest is dominated by the Guineo-Congolian Baikiaea insignis (eminii) and the Afromontane Podocarpus falcatus (P. usambarensis var. dawei). Other Guineo-Congolian species include: Canarium schweinfurthii, Klainedoxa gabonensis, Maesopsis eminii, Pseudospondias microcarpa, Pycnanthus angolensis, Symphonia globulifera and Tetrapleura tetraptera. T h e principal Afromontane tree species are: Apodytes dimidiata, Crotón megalocarpus, Ilex mitis, Podocarpus latifolius, Strombosia scheffleri, Suregada procera, Trichocladus ellipticus and Warburgia salutaris. T h e epiphytic lichen Usnea is abundant as in most types of Afromontane forest in East Africa. There has been heavy exploitation for timber, especially Podocarpus.
182
Vegetation of the floristic regions
Euphorbia dawei and Cynometra alexandri also occur in scrub forest in U g a n d a (T. J. Synnott, pers. c o m m . ) . Tall scrub forest (15 m tall) with an upper canopy of Strychnos potatorum, Tamarindus indica, Grewia mollis, Refs.: Germain (1952, p. 255-64); Lebrun (1947, p. 703-21; 1955, p. 59-64); Lewalle (1972, p. 57-69). Albizia grandibracteata, and Euphorbia candelabrum is Phots.: Germain (1952: 61); Lebrun (1947: 50, 51.1; 1955: 6). thought to represent the climax vegetation in the Ruzizi Profiles: Lewalle (1972: 14, 15). valley, though n o m o r e than tiny degraded vestiges Syn.: la forêt tropophile à Albizia grandibracteata et Strychnos remain. potatorum ['stuhlmannii'] (Germain, 1952); la forêt xérophile des crêtes: groupement à Crotón dichogamus et Evergreen and semi-evergreen bushland Euphorbia dawei (Lebrun, 1955); la forêt sclérophylle à Euphorbia dawei (Lewalle, 1972); la forêt sclérophylle à and thicket and derived communities Strychnos potatorum (Lewalle, 1972). (mapping unit 45) Scrub forest (mapping unit 45)
Vegetation intermediate between rain forest and evergreen and semi-evergreen bushland at one time probably occurred m o r e extensively in the Lake Victoria basin than in any other part of Africa, but few relics remain and published information is fragmentary. In U g a n d a and Burundi, Cynometra alexandri, which elsewhere is a canopy or emergent species of rain forest, also occurs in shorter forest and scrub forest and sometimes is less than 10 m tall. In scrub forest it is usually associated with Euphorbia dawei. In the basin of Lake Edward, Euphorbia dawei forms forest at 900-1000 m in bands u p to 3 k m wide along the banks of rivers and on the lower slopes of escarpments. T h e 12-15 m tall canopy is composed almost exclusively of this species and has a cover of 70-80 per cent. Since the trunks are w e a k and easily blown over, gaps in the canopy are numerous, but they are soon filled b y recruitment from young individuals, which regenerate freely in the shade of the parent trees. A few other species such as Euclea racemosa subsp. schimperi, Spathodea campanulata and Dombeya kirkii (mukole) are present in the canopy but only as isolated individuals. Lianes, especially Cissus quadrangularis, Bonamia paranoides, Senecio bojeri and Cissus petiolata, are numerous, and after reaching the canopy hang d o w n in draperies. Less vigorous lianes are Scutia myrtina and Cissus rotundifolia. Epiphytes are virtually absent except for the bracket fern Platycerium elephantotis. A n 8-10 m tall lower canopy is feebly developed beneath the Euphorbias but becomes dominant in the openings. It consists of Canthium vulgare, Cordia ovalis, Euclea racemosa subsp. schimperi and Olea africana. Other w o o d y species include Cassine aethiopica, Grewia similis, Carissa edulis, Erythrococca bongensis, Rhus natalensis and Teclea nobilis. T h e herb layer consists principally of Asystasia gangetica, Achyranthes áspera, Panicum deustum ana Justicia flava. A few mosses, including Fissidens sciophyllus, Racopilum speluncae and Archidium capense, are confined to the base of the trunks of Euphorbia dawei. In the Ruzizi valley, Euphorbia dawei forms scrub forest only in a single locality (Lewalle, 1972). Here it occurs as a 17-18 m tall emergent above a discontinuous 10-12 m tall canopy of Cynometra alexandri a n d Tamarindus indica. This particularfloristicassemblage is one of the most remarkable in the whole of Africa.
Refs.: Germain (1952, p. 233-51); Lebrun (1947, 2, p. 638- 61; 1955, p. 52-9); Lebrun & Gilbert (1954, p. 29-31, p.p.); Liben (1961); Troupin (1966). Phots.: Germain (1952: 55-9); Lebrun (1942: 36.2; 1947: 3942; 1955: 5); Liben (1961: 1). Profiles: Lebrun (1947: 95-6); Liben (1961: 1). Syn.: les forêts sclérophylles montagnardes et submontagnardes: ordre Oleo-Jasminetalia, alliance submontagnarde: Grewia-Carission edulis p.p. (Lebrun & Gilbert, 1954); les bosquets xérophiles à Maerua mildbraedii et Carissa edulis (Maerueto-Carissetum edulis); les bosquets xérophiles: association à Jasminum fluminense Çmauritianum') et Carissa edulis (Lebrun, 1955). Evergreen and semi-evergreen bushland and thicket, and related types of scrub forest (see above) probably represent the climax vegetation of large parts of this region. They have, however, been extensively destroyed, and today are represented b y small degraded relics which survive chiefly o n shallow soils, and by small patches of secondary regrowth. T o d a y the landscape is one of lightly wooded Acacia grassland with no m o r e than small islands of secondary evergreen thicket (mapping unit 45). Lebrun (1947, 1955) and Liben (1961) have suggested h o w patches of thicket b e c o m e established in secondary w o o d e d grassland dominated b y Acacia hockii, A. gerrardii, A. kirkii subsp. mildbraedii, A. Senegal and Euphorbia candelabrum. Lianes germinate in the shade of the Acacias and eventually smother their crowns. T h e shade they cast provides conditions suitable for the establishment of shrubs and bushes which, after a time, completely suppress the heliophilous Acacias which are unable to regenerate in the shade of the thicket. Euphorbia candelabrum (calycina) can also initiate the development of thicket. T h e shade it casts causes a diminution in the vigour of the grass layer which permits the invasion of hemi-heliophilous or even hemi-sciaphilous w o o d y plants. T h e principal species involved axe: Allophylus africanus, Azima tetracantha, Canthium schimperanum, Carissa edulis, Capparis fascicularis, C. tomentosa, Erythrococca bongensis, Grewia bicolor, Maerua triphylla {mildbraedii), Olea africana, Rhus natalensis, Tarenna graveolens and Turraea nilotica a m o n g bushy species, and Cissus quadrangularis, C. rotundifolia, Senecio stuhlmannii and Vernonia brachycalyx a m o n g climbers. T w o mosses, Bryum argenteum and Archidium capense, sometimes occur on the surface
The Lake Victoria regional mosaic
of the soil. Lock (1977¿>) suggests that Capparis tomentosa plays an important part in the establishment of thicket. T h e successional status of these thickets is uncertain. Lebrun originally (1947) thought that in the absence of
183
h u m a n interference they would be replaced by Euphorbia dawei scrub forest, but subsequently (1955) suggested that the latter is edaphically restricted, especially to rocky slopes, thereby implying that the thicket would be climax over extensive areas.
xiii The Zanzibar-Inhambane regional mosaic
Geographical position and area
Geographical position and area
Geology and physiography
This region occupies a coastal belt from southern Somalia (1°N.) to the m o u t h of the L i m p o p o River (25° S.). It is 50 to 200 k m wide except where it penetrates further inland along broad river valleys. Small exclaves also occur to the west on the windward slopes of mountainous massifs below 1500 m where the local increase in precipitation and dry-season relative humidity favours the development of lowland and transitional rain forest. (Area: 336000 k m 2 . )
Climate Flora Mapping units Vegetation Zanzibar-Inhambane lowland rain forest Transitional rain forest Zanzibar-Inhambane undifferentiated forest Zanzibar-Inhambane scrub forest Swamp forest Zanzibar-Inhambane transition woodland Zanzibar-Inhambane woodland and scrub woodland Zanzibar-Inhambane evergreen and semi-evergreen bushland and thicket Zanzibar-Inhambane edaphic grassland Zanzibar-Inhambane secondary grassland and wooded grassland
Geology and physiography Most of the land lies below 200 m but in the northern part there are scattered hills and plateaux rising considerably higher. They include the Shimba Hills (c. 400 m ) and M r i m a Hill in Kenya, the Pugu Hills and R o n d o (Mwera) Plateau in Tanzania, and the Macondes Plateau (986 m ) in northern Mozambique. Only the East U s a m b a r a M t s (1500 m ) in Tanzania exceed 1000 m . T h e outer part of the coastal belt, the coastal plain proper, is underlain by marine sediments of various ages from Cretaceous to recent, though in K e n y a there are also small areas of Jurassic outcrops. Inland from the coastal plain the more undulating topography is underlain principally by Precambrian rocks, but locally by Triassic sediments. T h e width of the coastal plain varies considerably and in northern M o z a m b i q u e is very narrow.
Climate Rainfall is mostly between 800 and 1200 m m per year and there is a well-defined dry season. Appreciably higher rainfall is experienced only in a few places such as the East U s a m b a r a M t s (Amani, 1946 m m ) and on the islands of Zanzibar and P e m b a (Wete, 1964 m m ) . In these places the amount and distribution is sufficient to support rain forest. In most parts of the Zanzibar-Inhambane Region the rainfall is comparable in amount to that of the Zambezian Region, but the dry season is less severe since relative humidity is high and n o m o n t h is absolutely dry. M e a n annual temperature is c. 26° C north of the Zambezi, but diminishes steadily southwards. Frosts are u n k n o w n . (See Fig. 18.)
The Zanzibar-Inhambane regional mosaic
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186
Vegetation of the floristic regions
Flora There are about 3000 species of which at least several hundred are endemic. Endemic genera. These include Cephalosphaera, Englerodendron, Grandidiera and Stuhlmannia. A further seven genera, including Bivinia, Hirtella, Ludia and Hymenaea (Trachylobium), which also occur in Madagascar, are confined or almost confined on the African mainland to this region. Hirtella and Hymenaea, although absent from West and Central Africa, are abundantly represented in the American tropics. Endemic species. T h e total n u m b e r is u n k n o w n but of the 190 forest tree species so far recorded, 92 (48.4 per cent) are endemic. T h e greatest concentration is in K e n y a and northern Tanzania, centred on the Shimba Hills and East U s a m b a r a M t s . Linking elements. O f the 190 forest trees species at present k n o w n , 15.3 per cent are Tongaland-Pondoland linking species, 4.7 per cent are Malagasy linking species, 25.8 per cent are Guineo-Congolian linking species, 3.7 per cent are Afromontane Unking species, and 7.0 per cent are ecological and chorological transgressors. Details of the Zambezian/ZanzibarInhambane linking element are given by Moll & White (1978).
Mapping units 16a. Z a n z i b a r - I n h a m b a n e coastal mosaic. 16b. Z a n z i b a r - I n h a m b a n e forest.
Vegetation Réf.: MoQSc White (1978). Forest is the m o s t widespread climax vegetation but has been largely replaced b y secondary w o o d e d grassland a n d cultivation. There are also extensive areas of scrub forest a n d edaphic grassland, a n d smaller areas of transition w o o d l a n d , bushland, a n d thicket. Typical w o o d l a n d does not occur in K e n y a , but from Tanzania southwards afloristicallyimpoverished type of m i o m b o w o o d l a n d b e c o m e s increasingly important. T h e forest is rich in species, but owing to the rapidity of change in a m o u n t a n d distribution of rainfall, dryseason atmospheric humidity, a n d the availability of soil moisture, it changes rapidly infloristiccomposition a n d p h y s i o g n o m y over quite short distances a n d is difficult to classify. O n l y the m o s t luxuriant types are classified as rain forest here. T h e latter is lowland rain forest except for that occurring o n the highest ground in the Z a n z i b a r - I n h a m b a n e Region, n a m e l y the summits of the m o s t elevated peaks in the East U s a m b a r a M t s ,
which are clothed with transitional rain forest. All forest other than rain forest is treated as a single continuum, n a m e l y undifferentiated Z a n z i b a r - I n h a m b a n e forest, m u c h of which has been referred to as rain forest b y previous authors. Zanzibar-Inhambane lowland rain forest (mapping units 16a and 16b) Refs.: Chapman & White (1970, p. 87, 97, 173); Pócs (19766, p. 478-9); Polhill (1968); White (MS, 1975-76); Wild & Barbosa (1967, p. 4). Phots.: Chapman & White (1970: 57). Profile: Chapman & White (1970: 7). Syn.: lowland rain forest (Tanzania, Polhill, 1968); lowland seasonal rain forest (Malawi, Chapman & White, 1970); moist evergreen forest at low and medium altitudes (Wild & Barbosa, 1967). Lowland rain forest was formerly extensively developed in Tanzania along the lower parts of the eastern highland arc, especially the Uluguru, Nguru, and Usambara M t s , and in parts of Ulanga and Iringa Districts. Only small fragments remain. Similar forest occurs on the lower slopes of mountains further inland, such as the Malawi Hills ( C h a p m a n & White, 1970), and thus forms small exclaves in the Zambezian Region. Most of the surviving forest in the East U s a m b a r a M t s occurs above 900 m and, because of its admixture of Afromontane species, is classified as lower transitional rain forest (see below). T h e main canopy of Zanzibar-Inhambane lowland rain forest, which is almost evergreen, is up to 20 m high, with émergents up to 40 m or more. This type differs from Guineo-Congolian rain forest in the greater degree of bud protection, less well-developed 'drip-tips' to the leaves, and the paucity of vascular and non-vascular epiphytes. T h e larger tree species include Aningeria pseudoracemosa (very local), Antiaris toxicaría, Burttdavya nyasica, Chlorophora excelsa, Khaya nyasica, Lovoa swynnertonii, Maranthes (Parinari) goetzeniana, Newtonia buchananii, Parkia filicoidea, Ricinodendron heudelotii, Steradia appendiculata a n d Terminalia sambesiaca. A small outlier of similar forest near Taveta in the S o m a l i a - M a s a i Region is s h o w n o n the m a p . It is d o m i nated b y Chlorophora excelsa, Cordyla africana, Diospyros mespiliformis a n d Newtonia buchananii, a n d o w e s its presence to the high water-table. Transitional rain forest ( m a p p i n g unit 16a) T h e rain forest near A m a n i in the East U s a m b a r a M t s in Tanzania is probably the best k n o w n in East Africa, at least to taxonomists, although it has never been fully described. T h e s u m m i t s of the East U s a m b a r a M t s (1254 m ) are not high e n o u g h for the occurrence of A f r o m o n t a n e rain forest, but several A f r o m o n t a n e species occur o n their upper slopes at altitudes appreciably lower than their n o r m a l lower limits o n
The Zanzibar-Inhambane regional mosaic
187
The fact that the East Usambara M t s harbour so other mountains. The forests in which they occur are m a n y species which are separated from their closest relathus transitional, though lowland elements predominate. tives by wide intervals suggests that they have served as The rainfall at A m a n i is 1946 m m per year. With a refugium for a formerly widespread flora which has decreasing altitude it falls off rapidly and rain forest become extinct over m u c h of its former area. probably does not descend below c. 800 m , although some species are found at lower altitudes in riparian Floristically poorer transitional rain forest also forest. occurs as small exclaves in the Zambezian Region, e.g. in Malawi at about 1370 m in the Misuku Hills, on Nchisi In relation to their small size the transitional rain M t , at Lisau Saddle and Chaone Hill in the Shire forests of the East Usambara M t s have a remarkably Highlands, and o n M a c h e m b a Hill near M t Mulanje rich and diversified flora. M o r e than 40 per cent of the (Chapman & White, 1970). T h e Chirinda forest in larger woody species are endemic to them or almost so, Z i m b a b w e (Banks, 1976) also belongs here. or, in a few cases, occur as endemic subspecies. T w o monotypic tree genera, Cephalosphaera (C. usambarensis) and Englerodendron (E. usambarense), are also Zanzibar-Inhambane endemic. Nearly 30 per cent of the larger woody species undifferentiated forest are Afromontane or, in East Africa at least, show upland (mapping units 16a and 16b) but not Afromontane distributions, and are characteristic members of lower transitional rain forest, though Refs.: Dale (1939, p. 8-11, 14-15); M o o m a w (1960, p. 22-9, none are confined to it. Most of the remaining species 35-40); White (MS, 1975-76). are Guineo-Congolian linking species. Phots.: Dale (1939: 2); M o o m a w (1960: 4, 7, 10). Syn.: lowland evergreen rain forest (Dale): Sterculia-ChloroThe majority of endemic species have their closest phora-Memecylon lowland rain forest (Moomaw); relatives in the lowland rain forests of the GuineoCongolian Region. S o m e species, e.g. Anonidium Combretum schumannii—Cassipourea lowland dry forest on usambarense, Enantia kummeriae, Isolona heinsenii and coral rag (Moomaw); evergreen dry forest (Dale); Polyceratocarpus scheffleri, belong to genera which are Manilkara-Diospyros lowland dry forest (Moomaw). otherwise confined to the Guineo-Congolian Region. A S o m e species are confined to the wetter types and some very wide interval separates the Usambara endemics to the drier, but m a n y occur throughout. Julbernardia from the other species in their respective genera. A magnistipulata, for instance, forms magnificant 30 m tall similar wide interval separates the endemic subspecies stands of moist forest but also occurs in scrub forest and (Greenwayodendron suaveolens subsp. usambaricum, evergreen bushland. Pterocarpus mildbraedii subsp. usambarensis and In the moister variants the main canopy occurs at Magnistipula butayei subsp. greenwayi) of those species 15-20 m and from it émergents rise to a height of 30 or which otherwise occur exclusively or principally in the 35 m . Very few individuals are taller than that. M a n y of Guineo-Congolian Region. the canopy species are briefly deciduous but not conThe Afromontane element is represented by Alancurrently so. Nevertheless this type is appreciably more gium chínense, Allanblackia stuhlmannii, Aningeria adolfi- deciduous than semi-evergreen lowland rain forest. friedericii, Cylicomorpha parviflora, Isoberlinia scheffleri, N o n e of the trees is buttressed, though some, e.g. Myrianthus holstii, Ocotea usambarensis, Strombosia Terminalia sambesiaca, areflutedat the base. Lianes are scheffleri, Syzygium sclerophyllum, Xymalos monospora plentiful but vascular epiphytes are usually scarce, as are and Zenkerella capparidacea s.l. bryophytes in general. Most tree trunks are without The species characteristic, inter alia, of lower transibryophytes except sometimes for a few minute hepatics. tional rain forest are: Macaranga capensis, Maranthes The richest forests are in Kenya and northern goetzeniana, Morinda asteroscepa, Newtonia buchananii, Tanzania. Towards the south there is progressive Strychnos mitis (mellodord) and Trichilia dregeana. floristic impoverishment and relatively few species reach Guineo-Congolian linking species include: AfroMozambique. sersalisia cerasifera, Antiaris toxicaría, Trilepisium The larger trees include: Afzelia quanzensis (20 m ) , madagascariense, Chrysophyllum perpulchrum, CleisAlbizia adianthifolia (25 m ) , Antiaris toxicaría (35 m ) , tanthus polystachyus, Ficus capensis, Funtumia africana, Apodytes dimidiata, Balanites wilsoniana (30 m ) , TrilePachystela msolo, Parkia filicoidea, Rauvolfia caffra, pisium madagascariense (20 m ) , Celtis wightii (20 m ) , Ricinodendron heudelotii, Schefflerodendron usambarense Cola clavata (20 m ) , Combretum schumannii (25 m ) , and Treculia africana. S o m e of these, e.g. Trilepisium, Cordyla africana (25 m ) , Chlorophora excelsa (35 m ) , Ficus capensis and Parkia, are widespread outside the Diospyros abyssinica (very rare), Diospyros mespiliformis Guineo-Congolian Region and occur in suitable (30 m ) , Erythrina sacleuxii (20 m ) , Erythrophleum habitats in the intervening region. Others, such as suaveolens (25 m), Fernandoa magnifica (20 m ) , Ficus Chrysophyllum perpulchrum, Pachystela msolo and Schefvallis-choudae (20 m ) , Inhambanella henriquesii (25 m ) , flerodendron usambarense, outside the GuineoJulbernardia magnistipulata (30 m ) , Lannea welwitschii Congolian Region are virtually confined to the (25 m ) , Lovoa swynnertonii (35 m ) , Macaranga capensis Usambara Mts. (25 m ) , Malacantha alnifolia (20 m ) , Manilkara sansibarensis (25 m ) , Mimusops aedificatoria (25 m ) , Newtonia
188
Vegetation ofthe floristic regions
paucijuga (25 m ) , Nesogordonia parvifolia (20 m ) , Paramacrolobium coeruleum (25 m ) , Parkia filicoidea (30 m ) , Pachystela brevipes (25 m ) , Rhodognaphalon schumannianum (30 m ) , Ricinodendron heudelotii (35 m ) , Sterculia appendiculata (35 m ) , Terminalia sambesiaca (35 m ) , Hymenaea verrucosa (30 m ) and Xylopia parviflora (holtzii) (25 m ) . T h e drier forests cover a larger area than the moister forests and extend further to the north and south. They are even more diverse floristically than the wetter forests and most of the larger tree species are sometimes gregarious and locally dominant or co-dominant. T h e principal trees are: Acacia robusta subsp. usambarensis (20 m ) , Afzelia quanzensis (15 m ) , Albizia petersiana (15 m ) , Balanites wilsoniana, Trilepisium madagascariense (15 m ) , Brachylaena huillensis (15 m ) , Cassipourea euryoides (15 m ) , Combretum schumannii (15 m ) , Cussonia zimmermannii (15 m ) , Cynometra webberi (12 m ) , Julbernardia magnistipulata (10-15 m ) , Manilkara sansibarensis (18 m ) , M . sulcata (10 m ) , Memecylon sansibaricum (9 m ) , Newtonia paucijuga (15 m ) , Oldfieldia somalensis (12 m ) , Pleurostylia africana (15 m ) , Scorodophloeus fischeri (15 m ) , Tamarindus indica (12 m ) and Hymenaea verrucosa (18 m ) . In m a n y places 25 m or more tall Chlorophora excelsa and Sterculia appendiculata are emergent. T h e cycad, Encephalartos hildebrandtii, and two succulent Euphorbias, E. nyikae and E. wakefieldii, are locally plentiful. Zanzibar-Inhambane scrub forest (mapping unit 16a) Réf.: White (MS, 1975-76). In K e n y a and southern Somalia scrub forest dominated by Diospyros cornii and Manilkara mochisia forms a quasi-continuous narrow band separating the forests of the coastal region from the bushlands of the interior. T h e annual rainfall is between 500 and 750 m m . Scrub forest reaches the coast between Malindi and L a m u where the rainfall is lower than elsewhere. Similar vegetation extends to southern Tanzania. D . cornii forms a discontinuous upper canopy 9-15 m tall. Sometimes the crowns are in contact but usually the cover is n o more than 50 per cent. A m o n g other canopy trees: Manilkara mochisia is an almost constant associate but is less plentiful; Dobera glabra is often abundant, especially where the water-table is near the surface; Newtonia erlangeri only occurs in the northern forests; and Terminalia spinosa indicates disturbance. Cactiform Euphorbias of tree dimensions, which are rare and often absent, are represented only by scattered individuals of E. candelabrum. T h e 1 m high E. grandicornis, however, frequently forms dense c o m munities in the understorey. T h e lower canopy, which reaches a height of 7 m , is very rich in species. It is usually dense but is only impenetrable where species of Sansevieria, Euphorbia grandicornis or the viciously spinous Adenia globosa
form dense thickets. T h e more characteristic members of this layer include: Bivinia jalbertii, Carissa sp., Crotón pseudopulchellus, Diospyros consolatae, Euclea natalensis, E. racemosa subsp. schimperi, Excoecaria venenífera, Grandidiera boivinii, Haplocoelum foliosum, H . inoploeum, Ochna thomasiana, Sideroxylon inerme, Suregada zanzibarensis, Thespesia danis, Thylachium africanum and Xeromphis nilotica. Climbers are rather rare, and epiphytes are virtually absent except between Witu and Garsen. T h e herb layer is poorly developed except for colonies of Sansevieria. M o s t species in scrub forest are evergreen. Only a few, e.g. species of Commiphora, are more than briefly deciduous. In most places D . cornii scrub forest has been degraded and converted into secondary deciduous bushland dominated by Albizia anthelminthica, Acacia bussei, A. mellifera, A. nilotica, Hyphaene compressa, Terminalia spinosa etc. S w a m p forest (mapping unit 16a) Fresh-water s w a m p forest is of restricted occurrence. Barringtonia racemosa forest with Acrostrichum aureum, Hibiscus tiliaceus, Pandanus spp. and Phoenix reclínala often occurs immediately behind the mangrove zone. In places Barringtonia extends upstream for a considerable distance. O n P e m b a Island Raphia s w a m p forest occurs in shallow valleys with very slow drainage. Associates include Elaeis guineensis, Voacanga thouarsii and the aroid Typhonodorum lindleyanum (Greenway, 1973). Zanzibar-Inhambane transition woodland (mapping unit 16a) Refs.: Dale (1939, p. 15-16); M o o m a w (1960, p. 30-5); White (MS, 1975-76); Wild & Barbosa (1967, p. 21, 26, 31). In places Zanzibar-Inhambane forest species occur in intimate mixture with heliophilous Zambezian w o o d land species to form communities which are intermediate between forest and woodland. S o m e such communities are clearly serai but others seem to be stable. In the Shimba Hills there are patches dominated by the woodland species Brachystegia spiciformis with an almost pure understorey of saplings of the forest species Paramacrolobium coeruleum. Near Witu in northern K e n y a the D o u m palm, Hyphaene compressa, is a conspicuous feature of the secondary grasslands. Locally the forest is encroaching and dead and dying Hyphaene have been overtopped by the forest species Trichilia emética, Erythrophleum suaveolens and Manilkara sansibarensis. Elsewhere in Kenya, as in the Arabuko-Sokoke forest, Brachystegia spiciformis forms almost pure stands on white sterile sands. T h e forest species Manilkara sansibarensis and Hymenaea verrucosa occur scattered in the canopy, and locally there are patches of evergreen thicket formed of forest shrubs and climbers. T h e forest
The Zanzibar-Inhambane regional mosaic
element, however, is not very vigorous and it appears that the soil is too unfavourable to permit the completion of the succession to forest. Zanzibar-Inhambane woodland and scrub woodland (mapping unit 16a) In a few dry rain-shadow areas, as in the foothills of the West Usambara Mts, the vegetation is a scrub woodland dominated chiefly by Zambezian linking species such as Cassia singueana, Combretum collinum, Dichrostachys cinérea, Heeria reticulata, Lonchocarpus bussei, Pappea capensis, Sclerocarya caffra, Stereospermum kunthianum and Ziziphus mucronata. The tallest species, Sclerocarya, is u p to 8 m high. Floristically poor m i o m b o woodland has a scattered distribution south of the R o v u m a River, where it interdigitates or occurs in mosaic with patches of forest and other vegetation. It forms parts of Wild & Barbosa's (1967) mapping units: 9, 10, 13, 25, 26, 27, 31, 32, and 33. Zanzibar-Inhambane evergreen and semi-evergreen bushland and thicket (mapping unit 16a)
189
height of 8 m . Associates include Syzygium cordatum, Uapaca sansibarica, Parinari curatellifolia, Manilkara sansibarensis, Euclea natalensis and scattered Pándanos goetzei. W h e n Philippia is exclusively dominant the ground is covered with leaf Utter or the lichen Cladonia medusiana; otherwise the fern Phymatodes scolopendria is conspicuous. Another species of Philippia, P. simii, forms an open shrubland on poorly drained soils o n the coast of Mozambique near Pebane (Wild & Barbosa, 1967). Zanzibar-Inhambane edaphic grassland (mapping unit 16a) In northern Kenya edaphic grassland studded with thicket-covered termite m o u n d s (see above) covers large areas of grey-black cracking clay soil near the mouth of the Tana River. It is largely treeless except for a few widely spaced individuals of Acacia zanzibarica, Hyphaene compressa, Terminalia spinosa and Thespesia danis. O n the coastal plain of Mozambique between the R . Sabi and the R . Buzi, the 'tandos' or seasonally flooded clayey depressions occurring o n sandy Quaternary or calcareous Cretaceous deposits are covered with Hyparrhenia, Ischaemum, Setaria grassland and are bordered by wooded grassland with Parinari curatellifolia, Uapaca nítida, Syzygiom guiñéense etc. There are also extensive areas of badly drained grassland on the deltas of the larger rivers.
Various types of bushland and thicket are found where unfavourable soil conditions prevent the development of forest. Dense thicket occurs on termite m o u n d s in seasonally waterlogged grassland which occupies parts of the coastal plain, as between Garsen and L a m u in northern Zanzibar-Inhambane secondary grassland Kenya. T h e thicket of Capparis, Carissa, Commiphora, and wooded grassland Euclea natalensis, Diospyros consolatae, Sideroxylon (mapping unit 16a) inerme etc. is usually overtopped by emergent trees of Diospyros cornii, Dobera glabra, Manilkara mochisia or These communities are extensive but published informaTamarindos indica. tion is meagre. In Kenya, between M o m b a s a and the According to Birch (1963) evergreen thicket repreTanzania frontier, the landscape is a mosaic of agrisents the climax o n shallow soils overlying coral limecultural crops, grassy fallows, secondary thicket often stone in parts of Kenya where the rainfall is between dominated b y Lantana, and orchards of Cocos, Anacar950 and 1200 m m per year. Characteristic species dium and Mangifera, which, w h e n the canopy is not too include Carpodiptera africana, Cussonia zimmermannii, dense, often have a carpet of grass. In places trees from Diospyros squarrosa, Zanthoxylum (fa*gara) chalybeum, the original forest, especially Chlorophora excelsa and Grewia plagiophylla, G. truncata, Haplocoelum inoploeum, Sterculia appendiculata, have been left standing. T h e Harrisonia abyssinica, Lannea stuhlmannii, Ludia mauri- palms, Borassus aethiopum and Hyphaene compressa, are tiana (sessiliflora), Manilkara sansibarensis, Millettia locally conspicuous. Other scattered non-forest trees usaramensis, Monanthotaxis fornicata, Pycnocoma littoinclude Acacia Senegal, Adansonia digitata, Afzelia quanralis, Sterculia rhynchocarpa, Suregada Zanzibarensis, zensis, Annona senegalensis, Antidesma venosum, CrosTabemaemontana elegans and Uvaria leptocladon. sopteryx febrífuga, Dalbergia melanoxylon, Dichrostachys cinérea, Flacourtia indica, Harrisonia abyssinica, Lannea A t a few places on the East African coast Ericaceous stuhlmannii, Lonchocarpus bussei, Maytenus senegalensis, bushland dominated by species of Philippia occurs o n Piliostigma thonningii, Sclerocarya caffra, Securidaca the waterlogged sites of former shallow lagoons or lake longependunculata, Stereospermum kunthianum, Strychbasins. Thus o n Mafia and P e m b a Islands (Greenway, nos madagascariensis, S. spinosa and Vitex mombassae. 1973), P. mafiensis forms a loose open canopy at a
xiv
T h e Kalahari-Highveld regional transition zone
Geographical position and area
Geographical position and area
Geology and physiography
T h e Kalahari-Highveld Transition Zone separates the Zambezian and K a r o o - N a m i b Regional Centres of E n d e m i s m . It runs diagonally across Africa from 13° S. in southern Angola to 33° S. in the Eastern Cape. Its width varies considerably. In the widest parts of the Highveld and Kalahari sectors it is more than 1800 k m across, but north of W i n d h o e k it suddenly narrows. (Area: 1223000 k m 2 . )
Climate Flora Mapping units Vegetation The Zambezia/Kaokoveld-Mossamedes transition Kalahari thornveld and the transition to Zambezian broad-leaved woodland The Windhoek Mountains The Kalahari/ Karoo-Namib transition Highveld grassland and associated communities Grassland Riparian scrub Rupicolous bushland and shrubland Scrub forest The Highveld/Karoo transition The transition from Afromontane scrub forest to Highveld grassland The Afromontane/Tongaland-Pondoland transition The Zambezia/Highveld transition
Geology and physiography Most of the Kalahari-Highveld transition zone occurs on the great Interior Plateau of southern Africa. In only a few places does it extend o n to or beyond the Great Escarpment. T h e Kalahari basin occupies the central part. F r o m this area of extremely low relief, which lies mostly between 850 and 1000 m , the landrisesgradually to the east and west. T h e peripheral highlands in the west, which are formed of Precambrian and Palaeozoic rocks, occupy only a narrow strip and reach their m a x i m u m elevation of 2484 m in the W i n d h o e k M t s . T o the east and south-east of the Kalahari basin the land rises gradually to more than 2000 m towards the plateaurimand the Great Escarpment, though for m u c h of its length the latter lies well within the Afromontane Region. The whole of this eastern part of the transition zone is underlain by Karoo rocks. T h e Kalahari basin is filled with sand and only locally are there outcrops of older rocks of Precambrian, Palaeozoic, and Karoo age. Its geological history has not been fully worked out, but it is thought that dune sands began to form at the end of the Cretaceous or in the early Tertiary. Since then there have been several changes of climate and the dunes in the extreme south m a y have assumed their present shape within the last 10000 years. Today the area is largely one of internal drainage but there have been periods of strongriverflow during the wetter phases of the Pleistocene. T h e K a a p Plateau, west of Kimberley, which is formed of Precambrian rocks and lies between 1220 and 1830 m , separates the Kalahari basin from the Karoo beds further east.
The Kalahari-Highveld regional transition zone
Climate Rainfall is intermediate between that of the Zambezian and K a r o o - N a m i b Regions. Nearly everywhere it is between 250 and 500 m m per year, but it increases somewhat in the east as the Drakensberg is approached. M o s t rain falls in summer. It is less concentrated than in the Zambezian Region but less evenly distributed than in the K a r o o - N a m i b . Winter temperatures are low everywhere except in the narrow north-eastern extension into Angola. Frost is widespread and severe, and absolute m i n i m u m temperatures are mostly lower than in the K a r o o N a m i b Region. (See Fig. 19.)
Flora
191
confined to the Kalahari-Highveld Region include Acacia haematoxylon, Anthephora argéntea and Schmidtia kalahariensis.
Mapping units 20. Transition from Afromontane scrub forest to Highveld grassland. 24. Mosaic of Afromontane scrub forest, Zambezian scrub woodland, and secondary grassland. 34. Transition from South African scrub woodland to Highveld grassland. 35a (p.p.). Transition from Zambezian undifferentiated woodland to Kalahari Acacia deciduous bushland and wooded grassland. 35c. T h e Windhoek Mountains. 36. Transition from Colophospermum mopane scrub woodland to K a r o o - N a m i b shrubland 44. Kalahari deciduous Acacia bushland and wooded grassland. 56. T h e Kalahari/Karoo-Namib transition. 57b. T h e Highveld/Karoo transition. 58. Highveld grassland.
T h e total flora is fairly large, possibly c. 3000 species, but this is because of the large number of marginal intruders which penetrate a short distance from the four contiguous major phytochoria. There are very few endemic species and the greater part of the interior has a very poor flora. Thus, Mostert (1958) records only 738 species of indigenous flowering plants from his study of the Bloemfontein and Brandford magisterial districts (2590 k m 2 ) . A high proportion of them, including Celtis africana, Commelina benghalensis, Crotalaria podocarpa, J uncus effusus, Phyllanthus maderaspatensis, SarcoThe Kalahari-Highveld Region abuts o n and provides stemma viminale, Tarchonanthus camphoratus, Themeda transitions connecting four major phytochoria. For this triandra and Typha australis, are pluriregional species. reason its vegetation pattern is complex. In the following S o m e 112 species (15.2 per cent) listed by Mostert are account the vegetation is not described primarily o n its Compositae and 111(15.1 per cent) are Gramineae. physiognomy as for other phytochoria, but the region is T h e flora of the southern Kalahari, which almost divided into nine subordinate areas, which largely exactly corresponds to mapping unit 56, is even poorer. coincide with the units shown o n the m a p and are Leistner (1967) records 438 species of flowering plants 2 described separately below. from the South African part (58000 k m ) , and estimates T h e greater part of the Kalahari-Highveld Region is that the total flora of the Southern Kalahari included in the Zambezian D o m a i n of the S u d a n o - Z a m (124320 k m 2 ) , which is appreciably larger than the entire bezian Region b y Werger (1978a), w h o does not C a p e floristic region, amounts to no more than recognize transition zones. H e places the remainder in 550 species. the K a r o o - N a m i b Region. Fewer than half the species Usted by Leistner can be regarded as typical K a r o o - N a m i b species. They include Lencosphaera bainesii, Nymania capensis, Parkinsonia africana, Phaeoptilum spinosum, Rhigozum trichotomum, The Zambezia/Kaokoveld-Mossamedes transition Stipagrostis amabilis and Tamarix usneoides. Most of the (mapping unit 36) remainder are widespread in southern Africa and m a n y Refs.: de Matos & de Sousa (1970); Giess (1971, p. 10); Tinley extend to the Zambezian Region or further, e.g. Acacia (1971); Volk (19666); Whellan (1965). erioloba, A. hebeclada, A. mellifera, Albizia anthelmintica, Boscia albitrunca, Diospyros lycioides, Terminalia sericea, Zambezian and Karoo-Namib species occur in intimate Echinochloa colona, Pogonarthria squarrosa and Sporomixture. Even Colophospermum mopane and Welwitschia bolns pyramidalis. O n e of the most characteristic grasses bainesii grow in the same community. A t its western of the Southern Kalahari, Asthenatherum glaucum, is limits in Angola at the edge of the Mossamedes desert, otherwise confined in South Africa to the K a r o o Colophospermum occurs as a scattered stunted tree 3 m N a m i b Region, but it also occurs near Lake Turkana in tall associated with the Zambezian species Acacia melliKenya. fera, Albizia anthelmintica, Commiphora sp., and Terminalia prunioides and abundant Welwitschia. T h e relatively few species which are more or less
Vegetation
192
Vegetation of the floristic regions
GHANZI (1131m) 207* 470 (20-84)
FIG. 19. Climate and topography of the Kalahari/Highveld regional transition zone (XIV) Figures 1-10 indicate the mapping units shown on the accompanying Vegetation Map of Africa as follows: (1) mapping unit 36, transition from Colophospermum mopane scrub woodland to Karoo-Namib shrubland; (2) mapping unit 44, Kalahari Acacia bushland and wooded grassland; (3) mapping unit 35c, the Windhoek Mts; (4) mapping unit 56, the Kalahari/Karoo-Namib transition; (5) mapping unit 58, Highveld grassland; (6) mapping unit 57b, the Highveld/Karoo transition; (7) mapping unit 20, transition from Afromontane scrub forest to Highveld grassland; (8) mapping unit 24, mosaic of Afromontane scrub forest, Zambezian scrub woodland and secondary grassland; (9) mapping unit 34, transition from South African scrub woodland to Highveld grassland; (10) mapping unit 35a, transition from Zambezian undifferentiated woodland to Kalahari Acacia deciduous bushland and wooded grassland. The stippled area shows the distribution of Kalahari Sand
The Kalahari-Highveld regional transition zone
Kalahari thornveld and the transition to Zambezian broad-leaved woodland (mapping units 35a, p.p., and 44) Refs.: Aco*cks (1975, p. 39-^3); Cole & Brown (1976); Wild & Barbosa (1967, p. 45, 46, 60). Phots.: Aco*cks (1975: 31, 32); Cole & Brown (1976: 7, 10); Volk (1966a: 3, 11); Walter (1971: 136, 137, 140, 141, 144-6).
193
Euclea crispa subsp. ovata, E. undulata, Euphorbia avasmontana, Grewia flava, Lebeckia macrantha, Maytenus heterophylla, Olea africana, Rhigozum obovatum, R. trichotomum, Rhus ciliata, R. dregeana, R. pyroides, R. lancea, R. undulata, Tarchonanthus minor and Ziziphus mucronata. T h e W i n d h o e k Mountains (mapping unit 35c)
W o o d e d grassland is the characteristic vegetation of the Refs.: Giess (1971, p. 11); Volk & Leippert (1971). thick mantle of Kalahari Sand. In Botswana the m o r e Phots.: Giess (1971: 39-42). or-less continuous grass sward is less than 1 m high and Giess's photographs s h o w a w o o d e d grassland, but the consists principally of Anthephora argéntea, A . pubescens, Digitaria pentzii, Eragrostis biflora, E. ciliaris, E. original vegetation w a s probably denser. T h e flora is a mixture of Zambezian and K a r o o - N a m i b species. T h e lehmanniana, E. pallens, Panicum kalaharense, P. lanipes, principal trees and bushes are Acacia hereroensis, Pogonarthria squarrosa, Schmidtia kalahariensis, S. papCombretum apiculatum, Acacia reficiens subsp. reficiens, pophoroides and Stipagrostis uniplumis. O f these, AntheA . hebeclada, Euclea undulata, Dombeya rotundifolia, phora argéntea in almost endemic, and a few species, e.g. Tarchonanthus camphoratus, Rhus marlothii, Albizia Panicum lanipes and Schmidtia kalahariensis, are shared anthelmintica, Heeria (Ozoroa) crassinervia, Ficus cordata with the K a r o o - N a m i b Region, but the majority extend and F. guerichiana. T h e original grass cover consisted from the K a r o o - N a m i b at least as far as the southern of Anthephora pubescens, Brachiaria nigropedata, part of the Zambezian Region. Cymbopogon spp., Heteropogon contortus, Hyparrhenia T h e principal trees and bushes in the southern hirta, and others, but these grasses are n o w sparse in variant (mapping unit 44) are all Zambezian species, m a n y parts because of overgrazing. T h e K a r o o - N a m i b namely Acacia erioloba, A . fleckii, A . hebeclada, A . luederitzii, A . mellifera, A . tortilis, Boscia albitrunca, Dichro- grasses, Stipagrostis obtusa, S. uniplumis, and Panicum lanipes, also occur. Phaeoptilum spinosum and species of stachys cinérea and Terminalia sericea. In the northern Aptosimum, Eriocephalus, Galenia, Pentzia, Plinthus, variant (mapping unit 35a) broad-leaved trees are m o r e Salsola and Tetragonia represent the K a r o o - N a m i b abundant, and include Combretum collinum, Commiflora a m o n g small shrubs and forbs. phora africana, C. angolensis, Ochna pulchra and Ziziphus mucronata, but Acacia is still dominant. T h e trees are always less than 7 m tall and usually are m u c h smaller. Normally they are widely spaced. O n the shallow soils of the quartzite and limestone outcrops which form the Ghanzi Ridge the w o o d y plants are denser, grasses are relatively less important and shrubs are plentiful. T h e latter include the K a r o o species Rhigozum brevispinosum, Leucosphaera bainesii, Phaeoptilum spinosum and Montinia caryophyllacea. In the C a p e Province north of the Orange River, Kalahari Sand alternates with extensive areas of stony soil o n the K a a p Plateau, Langeberg, and Asbestos mountains. O n Kalahari Sand in this area the vegetation has been severely degraded. Themeda triandra w a s formerly the dominant grass but it has been extensively replaced, because of overgrazing, b y 'white' desert grasses, chiefly species of Aristida, Eragrostis and Stipagrostis. Further degradation leads to a uniform sward of Schmidtia pappophoroides, and ultimately to invasion by the K a r o o shrublets Pentzia incana and Chrysocoma tenuifolia. T h e most abundant and characteristic tree, Acacia erioloba, has been removed from large areas to provide fuel for the mines at Kimberley. O n the most stony soils the vegetation is dense bushland. T h e principal species is Tarchonanthus camphoratus. Its w o o d y associates include: Acacia karroo, A. mellifera, A . tortilis, Boscia albitrunca, Buddleja saligna, Crotón gratissimus, Diospyros lycioides, Ehretia rígida,
T h e Kalahari/Karoo-Namib transition (mapping unit 56) Refs.: Coetzee & Werger (1975, p. 549-50); Giess (1971, p. 13); Leistner (1967); Leistner & Werger (1973); Walter (1971, p. 256-8). Phots.: Coetzee & Werger (1975: 20); Giess (1971: 65-7); Leistner (1967: 1, 2, 8, 18, 36, 39-42); Leistner & Werger (1973:2,4). Profile: Walter (1971: 148). This is an area of wind-blown sand which occurs as fixed dunes in the form of long parallel ridges. Sand covers 9 0 per cent of the surface. In undisturbed areas, the lower slopes of dunes are largely consolidated by vegetation but the upper slopes and crests are subjected to erosion b y strong winds and the cover is m u c h sparser. T h e vegetation is a mosaic of lightly w o o d e d grassland o n the dune crests, pure grassland in shallow depressions between the dunes, and Rhigozum trichotomum shrubby grassland in deeper hollows where the underlying calcrete is near the surface. In undisturbed places the grasses are mostly perennials including Asthenatherum glaucum, Stipagrostis uniplumis, Eragrostis lehmanniana, Stipagrostis ciliata, and, o n the dune crests, Stipagrostis amabilis. A c o m m o n pioneer is Megaloprotachne albescens, whereas in disturbed areas in drier regions the annual Schmidtia kalahariensis is dominant. T h e c o m m o n e s t trees are Acacia
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Vegetation of the floristic regions
erioloba, Boscia albitrunca, Acacia reficiens subsp. reficiens, Albizia anthelmintica a n d Terminalia sericea, w h i c h often occurs as a shrub. Acacia haematoxylon, w h i c h is almost confined to this region, is usually a shrub, but sometimes a tree.
Highveld. It usually forms a sward 25-75 c m tall, which in s u m m e r looks dense, but the basal cover rarely exceeds 25 per cent. In winter and during droughts the grass cover is m u c h shorter and the associated forbs and bare spaces are m o r e conspicuous. It has been widely assumed that Themeda is the natural climax species throughout most of the Highveld Highveld grassland and associated communities area, but there is some evidence (Roux, 1969) that its (mapping unit 58) dominance depends on fire, and that with fire-protection Themeda is partly replaced b y other species. In the Refs.: Aco*cks (1975, p. 87-95); Coetzee & Werger (1975, wetter parts of the Highveld, Themeda is commonly p. 551-3); Mostert (1958, p . 85-161); V a n Zinderen associated with Elionurus argenteus, Heteropogon conBakker, Jr(1971, 1973); Werger (1973a, p. 113-27). tortus, Trachypogon spicatus, and Tristachya leucothrix. Phots.: Aco*cks (1957: 81-90); Coetzee & Werger (1975: 22); Van Zinderen Bakker, Jr (1973: 1). In drier types the lower-growing species, Aristida congesta, Eragrostis lehmanniana and Tragus bertëroHighveld grassland represents the climatic climax nianus, are plentiful. between 1220 and 2150 m o n large parts of the high O v e r extensive areas, because of overgrazing, interior plateau in South Africa which extends west of Themeda h a s b e e n largely eliminated a n d h a s been the Drakensberg from the extreme south of the Transreplaced b y pioneer grasses such as Aristida spp. a n d vaal through the Orange Free State to the Eastern Cape. Chloris virgata as well as invasive K a r o o shrublets T h e total tree flora of this region is very small and the (Chrysocoma tenuifolia) a n d annual w e e d s (Tribulus development of w o o d y vegetation is precluded nearly terres tis). everywhere b y the dry, extremely frosty winters. T h e following forbs are of general occurrence in o n e Although fire is almost certainly a natural ecological or m o r e of A c o c k s ' s types: Ajuga ophrydis, Anthofactor there is insufficient evidence that it is primarily spermum rigidum, Asclepias multicaulis, Barleria responsible for the almost total absence of larger w o o d y macrostegia, Berkheya onopordifolia, B . rígida, Conyza plants. T h e latter are virtually confined to riparian pinnata, Crabbea acaulis, Cyperus obtusiflorus, Dicoma forest, bushland, and thicket on the few rocky hills and macrocephala, Euphorbia inaequilatera, E . striata, Felicia escarpments, and to scrub forest in sheltered ravines in filifolia, F. muricata, Geigeria áspera, Gnidia kraussiana, the foothills of the Maluti M t s and the Drakensberg in Haplocarpha scaposa, Helichrysum dregeanum, H . latithe east. folium, H . rugulosum, H . oreophilum, Hermannia betonicifolia, H . coccocarpa, H . depressa, Hypoxis rigidula, H . rooperi, Indigofera alternons, I. rostrata, Ipomoea crasGrassland sipes, Kohautia amatymbica, Osteospermum scariosum, Oxalis depressa, Rhynchosia totta, Scabiosa columbaria, Aco*cks (1975) recognizes 10 types of Highveld grassScilla nervosa, Senecio coronatus, S. erubescens, Sonchus land, which are distinguished mainly by the different nanus, Stachys spathulata, Vernonia oligocephala, Walaproportions in which a handful of species occur. T h e frida densiflora a n d W. saxatilis. Ziziphus zeyherana is a following species are of general occurrence in one or rhizomatous geoxylic suffrutex. more of his types: Alloteropsis semialata, Andropogon Several of the above belong to genera, e.g. Berkheya, amplectens, A. appendiculatus, A. schirensis, Anthephora Geigeria, Gnidia, Haplocarpha, Helichrysum, Osteopubescens, Aristida congesta, A. junciformis, Brachiaria Oxalis, a n d Walafrida, w h i c h have their serrata, Chloris virgata, Ctenium concinnum, Cymbopogon spermum, plurinodis, Cynodon dactylon, C. incompletes, Digitaria greatest concentration of species in South Africa. T h e argyrograpta, D. diagonalis, D. monodactyla, D. tricho- majority, h o w e v e r , belong to predominantly tropical or laenoides, Elionurus argenteus, Eragrostis atherstonei, E. subcosmopolitan genera. capensis, E. chloromelas, E. gummiflua, E. lehmanniana, E. micrantha, E. obtusa, E. plana, E. racemosa (chalRiparian scrub cantha), E. sclerantha, E. superba, Eustachys paspaloides, Harpechloa falx, Heteropogon contortus, Microchloa Principal species are Acacia karoo (7 m ) , Celtis africana, caffra, Monocymbium ceresiiforme, Panicum coloratum, Diospyros lycioides, Rhus lancea a n d Ziziphus mucronata. P. natalense, Pogonarthria squarrosa, Setaria flabellata, S. nigrirostris, S. sphacelata, Sporobolus discosporus, S. Rupicolous bushland and shrubland fimbriatus, Themeda triandra, Trachypogon spicatus, Tragus koelerioides, T. racemosus, Trichoneura grandiIn the drier lower-lying western parts the following glumis, Triraphis andropogonoides and Tristachya leuco- bushes are characteristic: Acacia karroo, Buddleja thrix {hispida). T h e absence from this list of Hyparsaligna, Celtis africana, Cussonia paniculata, C. spicata, rhenia, the tall species of which are so conspicuous in the Diospyros austro-africana, D . lycioides, Ehretia rigida, Zambezian Region, is noteworthy. Themeda triandra is Euclea crispa, Grewia occidentalis, Heteromorpha arborby far the most widespread and abundant species in the escens, Olea africana, Osyris sp., Rhus ciliata, R . erosa, R .
The Kalahari-Highveld regional transition zone
lancea, R. undulata, Tarchonanthus camphoratus, and Ziziphus mucronata. Towards the east, Afromontane species b e c o m e increasingly prominent. In the Kimberley area there are karroid communities characterized b y Chrysocoma tenuifolia, Cotyledon decussata, Eberlanzia spinosa, Eriocephalus spinescens, Euphorbia mauritanica, Pentzia sphaerocephala, Rhigozum obovatum and Ruschia unidens. F r o m about Maseru southwards Aloe ferox is conspicuous o n northern slopes. Even the most luxuriant communities are rarely m o r e than 5 m tall.
Scrub forest
195
Because of overgrazing the grassland has been converted to a secondary K a r o o dwarf shrubland similar to that of the Central U p p e r K a r o o but m o r e grassy and floristically poorer. T h e principal surviving grasses are Aristida congesta, Cynodon hirsutus, Eragrostis curvula, E. lehmanniana, E. obtusa, Themeda triandra and Tragus koelerioides. T h e most abundant K a r o o shrublets include Chrysocoma tenuifolia, Aptosimum procumbens (depressum), Gnidia polycephala, Hermannia conocarpa, Pentzia globosa and Walafrida saxatilis. This type occurs on the rather deep soils of the pediplains. B y contrast, the lithosols of mesas, kopjes, and ridges support various types of shrubland from which the shrublets of the false K a r o o have been recruited, and, m u c h m o r e locally, bushland. T h e principal species of these shrubland and bushland communities are:
Patches of 10 m tall scrub forest composed predominantly of Afromontane species occur between 1500 Large shrubs and bushes: Buddleja saligna, Celtis afriand 1900 m in the eastern parts of the Orange Free State cana, Cussonia paniculata, Diospyros austro-africana, D . and adjacent parts of Lesotho. They are confined to lycioides, Ehretia rigida, Euclea crispa, subsp. ovata, shallow water-retaining soils overlying consolidated Maytenus polyacantha, Olea africana, Osyris sp., Rhiscrees in deep ravines. Because of heavy exploitation for gozum obovatum, Rhus ciliata, R. erosa, R. undulata, firewood and degradation by cattle only small fragments Tarchonanthus camphoratus and Ziziphus mucronata. remain. T h e principal species are: Buddleja salviifolia, Cassinopsis ilicifolia, Celtis africana, Diospyros whyteana, D w a r f shrubs: Aptosimum procumbens, Eriocephalus Euclea coriácea, E. crispa, Grewia occidentalis, Halleria spinescens, Hermannia candidissima, Hibiscus marlolucida, Ilex mitis, Kiggelaria africana, Leucosidea sericea, thianus, Pegolettia retrofracta and Pentzia sphaerocephala Maytenus acuminata, M . heterophylla, M . undata, as well as the ubiquitous Chrysocoma tenuifolia and Myrsine africana, Olea africana, Olinia emarginata, Pentzia globosa. Osyris sp., Pittosporum viridiflorum, Podocarpus latifolius, Rhamnus prinoides, Rhus pyroides and Scolopia mundii. Grasses: Aristida diffusa, Enneapogon desvauxii, FingerO f these, Celtis is deciduous. Leucosidea, Kiggelaria and huthia africana, Heteropogon contortus, Hyparrhenia hirta . Maytenus acuminata are semi-deciduous. T h e others are and Themeda triandra. evergreen.
T h e Highveld/Karoo transition (mapping unit 57b) Refs.: Aco*cks (1975, p. 78-81); Werger (1973a, 19736). Phots.: Aco*cks (1975: 72, 73, 75, 76); Werger (1973b: 2-4). Syn.: false Upper Karoo; false karroid broken veld, false Central Lower Karoo; pan turf veld invaded by Karoo; karroid Merxmuellera mountain veld replaced by Karoo (all of Aco*cks). Apart from riparian scrub forest, and various types of shrubland and bushland on rocky slopes, the whole region is believed to have formerly been grassland similar to the surviving Highveld grassland of today. Early travellers (summarized b y Werger, 1973a) describe the general grassiness of the region and the absence of shrubs and bushes, besides commenting o n the multitude of wild animals. It appears that at the end of the eighteenth century, the country near the present Colesberg, which w a s then outside the borders of the C a p e Colony, had a n abundance of grass, but that thirty-five years later the grass had largely been replaced by dwarf shrubs. In the area north of the Orange River, not at that time included in the colonized areas, grassland w a s still the dominant vegetation type.
Succulents: Euphorbia clavarioides, Haworthia tesselata and Pachypodium succulentum. T h e deep sandy levees fringing the Orange River support 6-10 m tallriparianforest composed principally of Acacia karroo, Celtis africana and Diospyros lycioides.
T h e transition from Afromontane scrub forest to Highveld grassland (mapping unit 20) Refs.: Jacot Guillarmod (1971); Killick (1978c, p. 540-2). Phot.: Killick (1978c: 10). Published information is meagre. Although the Natal slopes of the Drakensberg are occupied b y Afromontane communities between the 1280 m contour and the summit area, the corresponding slopes in Lesotho below about 2900 m are almost bereft of the most typical Afromontane elements. H e n c e this area must be excluded from the Afromontane Region. T h e Lesotho slopes are almost entirely covered by Themeda-Festuca grassland. There are only isolated patches of scrub dominated by Leucosidea sericea, Buddleja corrugata, Passerina montana and species of Erica. In places Leucosidea is
196
Vegetation of the floristic regions
less than 2 m tall and is closely pressed to the ground. Dalbergia obovata, Ekebergia pterophylla, Ficus capensis, O n northern slopes Themeda prevails u p to 2750 m , F. sonden and Syzygium cor datum. above which Festuca caprina becomes dominant. T h e T h e most abundant grasses are Andropogon schirengrasses, however, are often replaced by Chrysocoma sis, Brachiaria serrata, Elionurus argenteus, Eragrostis tenuifolia or Felicia filifolia as a result of overgrazing. O n racemosa, Heteropogon contortus, Hyparrhenia hirta, southern slopes Festuca caprina descends as a dominant Monocymbium ceresiiforme, Rendlia altera, Themeda to 2135 m . triandra, Trachypogon spicatus and Tristachya leucothrix. T h e Afromontane/Tongaland-Pondoland transition (mapping unit 24)
T h e Zambezia/Highveld transition (mapping unit 34)
Réf.: Aco*cks (1975, p. 100-3). Phots.: Aco*cks (1975: 97-9).
Refs.: Aco*cks (1975, p. 99); White (1978a, p. 477-9). Phots.: Aco*cks (1975: 96).
This transition separates the Tongaland-Pondoland Region from the Afromontane vegetation of the Natal Drakensberg north of the Tugela basin, and continues o n the upper slopes of the ridge which connects the Natal and Transvaal sectors of the Drakensberg. It lies chiefly between 800 and 1700 m . T h e vegetation today is chiefly grassland but originally w a s probably bushland with scrub forest in sheltered kloofs. T h e w o o d y relics are chiefly Afromontane at higher altitudes and include Apodytes dimidiata, Halleria lucida, Leucosidea sericea, Pittosporum viridiflorum, Podocarpus latifolius, Rapanea melanophloeos and Scolopia mundii. 'Lowland' w o o d y species include Acacia caffra, A. davyi, A. nilotica subsp. kraussiana, A. sieberana, Aloe arborescens, Celtis africana, Commiphora harveyi,
This area coincides with parts of Aco*cks's veld type 61, Bankenveld. Its original vegetation w a s probably bushland dominated by Acacia caffra, but the prevalent vegetation today is secondary grassland. There are relatively few tree species and their n u m b e r diminishes rapidly towards the south. Afromontane and Zambezian species are present, surviving chiefly in bushland and scrub forest in sheltered kloofs. Afromontane species include Calodendrum capense, Diospyros whyteana, Halleria lucida, Ilex mitis, Kiggelaria africana, Leucosidea sericea, Nuxia congesta, Olinia and Pterocelastrus. Zambezian species include Acacia robusta, A. sieberana, Burkea africana, Combretum molle, Dombeya rotundifolia, Ficus ingens, F. soldanella, Lannea discolor, Mimusops zeyheri, Ochnapulchra and Strychnos pungens.
xv
The Tongaland-Pondoland regional mosaic
Geographical position and area
Geographical position and area
Geology and physiography
This Region extends from the L i m p o p o River mouth (25° S.) to Port Elizabeth (34° S.). In the north it is up to 240 k m wide, but locally in the south where mountains c o m e close to the sea its width is no more than 8 k m . Elsewhere in the south it penetrates inland along river valleys far into the interior. For most of its length it lies below the Afromontane Region or the Afromontane/ Tongaland-Pondoland transition zone (page 196). (Area: 148000 k m 2 . )
Climate Flora Mapping units Vegetation Tongaland-Pondoland undifferentiated forest Tongaland-Pondoland scrub forest Tongaland-Pondoland swamp forest Tongaland-Pondoland evergreen and semi-evergreen bushland and thicket Tongaland-Pondoland edaphic grassland Tongaland-Pondoland secondary grassland
Geology and physiography T h e coastal plain in the north is composed of Cretaceous and Tertiary marine sediments. Elsewhere, the more undulating landscape which rises locally to 1600 m is carved out of rocks of the Basem*nt Complex, Table Mountain Sandstone and sedimentary strata of the Karoo system.
Climate O w i n g to the ameliorating effect of the w a r m M o z a m b i q u e Current the coastal regions have a moderately high and well-distributed rainfall and, except in the extreme south, are frost-free. Further inland, however, climate changes rapidly over short distances and there is often a great contrast between xerocline and mesocline vegetation. T h e desiccating 'berg' winds have a profound effect on valley vegetation. T h e rainfall is more evenly distributed throughout the year than in most parts of the Zanzibar-Inhambane Region. M e a n annual temperature diminishes from 22° C in the north to 17° C in the south. (See Fig. 20.)
Flora There are about 3000 species. M o r e than 200 of the larger w o o d y species, approximately 40 per cent of the total, are endemic. T h e proportion of endemic herbs and smaller w o o d y plants is u n k n o w n but is probably less. Endemic family. T h e Achariaceae is centred on this region, but is not strictly endemic.
198
Vegetation of the floristic regions
BARBERTON (152m) 20-3* 777 (20-54) HLABISA (245m) It-»* 1121 (13-30)
FIG. 20. Climate and topography of the Tongaland-Pondoland regional mosaic ( X V )
MAPUTO (35)
S»m) 22-2'003
The Tongaland-Pondoland regional mosaic
Endemic genera. T h e twenty or so endemic w o o d y genera include Anastrabe, Bachmannia, Burchellia, Ephippiocarpa, Galpinia, Harpephyllum, Hippobromus, Jubaeopsis, Loxostylis, Pseudosalacia, Rhynchocalyx, Stangeria a n d Umtiza. T w o genera, Atalaya a n d Protorhus, occur n o w h e r e else in Africa but are found in Asia a n d Madagascar. Other characteristic genera. Twenty-six of the 35 South African species of Encephalartos, 12 of the 23 South African tree Aloes and nine of the 13 South African succulent tree Euphorbias occur in Tongaland-Pondoland. Linking elements. Of the 500 or so larger woody plants occurring in Tongaland-Pondoland, 7.6 per cent are Zanzibar-Inhambane linking species, 20 per cent are Zambezian Unking species, 8.7 per cent are Afromontane linking species, 5.1 per cent are GuineoCongolian linking species, and 2.5 per cent are KarooN a m i b linking species. A further 1.5 per cent also occur in upland areas in tropical Africa but are not truly Afromontane. The importance of the Zambezian element decreases and that of the Karoo-Namib element increases towards the south. The representation of the Afromontane element is higher than in any other lowland phytochorion (Moll & White, 1978).
Mapping units 16c. T h e Tongaland-Pondoland coastal mosaic. 29e. T h e transition from undifferentiated Z a m b e z i a n w o o d l a n d to Tongaland-Pondoland bushland (see page 96 a n d below). 39. Tongaland-Pondoland evergreen a n d semi-evergreen bushland a n d thicket. 48. Tugela basin w o o d e d bushland.
Vegetation Where the vegetation has not been completely destroyed it consists of a complex mosaic of forest, scrub forest, and evergreen and semi-evergreen bushland and thicket in a matrix of secondary grassland and wooded grassland. There are small patches of woodland in the north and of edaphic grassland and swamp forest on the coastal plain. The vegetation has been described in some detail by Moll & White (1978). Tongaland-Pondoland undifferentiated forest (mapping unit 16c) Refs.: Aco*cks (1952, 1975: mapping units 1 p.p., 2, 6 p.p.); Breen (1971); Edwards (1967, p. 82-6); Huntley (1965); Killick (1958, p. 60-72); Moll (19686, 1968res
*
Madeira "¥•
Canary Is. 4.'^.
*
20' C a p e Verde Is.
*
Ascension^
St.Helena*
20'
Tristan da Cunha If0 1
500 •
1000 Km 1
F I G . 24. Islands of the eastern Atlantic O c e a n
245
246
Vegetation of the floristic regions
Canary islands as continental fragments with volcanic overlays. There is a wide range of climates. T h e Azores are colder and wetter than the more southerly archipelagos. B y comparison, Madeira is m u c h warmer and, although there is a north-south gradient in rainfall, the vegetation zones are relatively uniform all around the island. T h e climate of the Canaries, by contrast, shows m u c h wider extremes and this is reflected in its richer flora and m o r e diverse vegetation. Cloud belts are restricted to northerly or north-easterly slopes, while the southern slopes are m o r e arid. Because of the high altitude of some islands several altitudinal zones of vegetation can be recognized. T h e tropical Cape Verde Islands are uniformly arid except in the mountains and show only slight zonation. Throughout Macaronesia the vegetation has been severely degraded by m a n , and several endemic species are believed to be threatened with extinction (Lucas & Synge, 1978). Although there are m a n y publications dealing with the floristics and plant taxonomy of Macaronesia, detailed ecological studies are few and there are n o syntheses. T h e total flora of Macaronesia comprises approximately 3200 species of flowering plants, of which c. 680 (20 per cent) are said to be endemic (Humphries, 1979). Included in the former total, however, is a large number of introduced species, so that in fact the endemics are proportionally more numerous. Generic endemism is relatively weak, amounting to only 31 (12.4 per cent) out of a total of 251 genera. Eighteen are confined to a single archipelago (17 in the Canaries, one in Madeira). T h e majority of endemic genera are m o n o - or oligospecific, only four (Aichryson, Argyranthemum, Monanthes, Sinapidendrori) having more than five species. Only three endemic genera {Picconia, Pleiomeris and Visnea) include large trees. M o s t of the non-endemic genera are also poor in species, but a few genera such as Aeonium (36 species), Sonchus (29 species), Echium (28 species), Lotus (27 species) and Argyranthemum (endemic, 22 species) have undergone a remarkable adaptive radiation, especially in the Canary Islands. A distinctive feature of the Macaronesian flora is the large n u m b e r of arborescent, often pachycaul, species in otherwise predominantly herbaceous genera, such as Echium, Sonchus, Limonium, Plantago and Sanguisorba. M o s t of the tree species are endemic, notably Apollonius barbujana, Arbutus canariensis, Clethra arbórea, Cytisus (Teline) stenopetalus, Dracaena draco, Erica scoparia subsp. azorica, Euphorbia tuckeyana, Heberdenia excelsa (bahamensis), Ilex canariensis, I. perado, I. perado subsp. platyphylla, Juniperus brevifolia, J. cedrus, Laurus azorica, Ocotea foetens, Persea indica, Phoenix atlántica, P. canariensis, Picconia (Notelaea) azorica, P. excelsa, Pinus canariensis, Pittosporum coriaceum, Pleiomeris canariensis, Sideroxylon marmulano and Visnea mocanera. T h e floristic relationships of Macaronesia are extremely diverse, including affinities with America. T h e
largest elements, however, consist of Mediterranean linking species, and of endemic species of Mediterranean affinity. T h e floras of the arid lowlands in the Canaries and C a p e Verde Islands are closely related to those of the nearby African mainland. Other linking elements involving more distant parts of Africa include: 1. Erica arbórea: Canaries, Madeira, Mediterranean Region, Sahara M t s (Tibesti), M t s of E . Africa from Ethiopia to southern Tanzania. 2. Myrsine africana: Azores, African mainland from the R e d Sea Hills southwards to the Cape, westwards from Tanzania to Angola, and eastwards to China. 3. Canarina canariensis: Canaries; C. abyssinica and C. eminii o n E . African mountains from Ethiopia to Tanzania. 4. Ocotea foetens: Madeira, Canaries; O. gabonensis, G a b o n , C o n g o Republic; O. bullata (including O. kenyensis), Ethiopia to the Cape. 5. Dracaena draco: Canaries, Cape Verde Islands. D . ombet: Egypt south to Ethiopia. D . cinnabari: Socotra. 6. Visnea mocanera: Canaries. Balthasaria mannii: Sâo T o m é . B. schliebenii: East African Mts. There can be little doubt that the history of the Macaronesian flora is complex, and m a n y unresolved problems remain. It is widely believed that the 'Laurel' forest represents a relic of a humid subtropical flora which was widespread in southern Europe and parts of North Africa during the Upper Tertiary. T h e Azores Refs.: Dansereau (1966); Guppy (1917); Marier & Boatman (1952); Sjogren (1973); Tutin (1953). T h e Azores are a group of nine islands and a number of rocks lying between about 37° and 39° N . and 25° and 32° W . Their total land area is about 1800 k m 2 . T h e island of Fayal in the middle of the group is 450 k m from Lisbon and 1900 k m from Newfoundland. T h e islands are all volcanic and of recent origin. Pico, the highest island,risesto 2300 m . T h e climate is extremely oceanic, characterized by a moderate rainfall spread evenly throughout the year, high relative humidity, and a small temperature range. Frost occurs at high elevations. O f the 700 or so species of flowering plants at least 200 have been introduced. About 40 are endemic. There are n o endemic genera. T h e affinity of the flora is overwhelmingly European, though the trees of the 'Laurel forest' are all, with the exception of Myrica faya and Persea indica, endemic species or varieties, nearly all of which appear to be closely allied to Madeiran species. /. Coastal vegetation T h e most abundant species include Solidago sempervirens, Juncus acutus, Euphorbia azorica and the grasses Cynodon dactylon, Agrostis azorica and Polypogon monspeliensis. A lava flow o n the eastern side of Pico
Other offshore islands
produced by an eruption in 1718 which was still uncultivated in 1929 gave an indication of the natural vegetation. My rica faya (2-3 m high) was dominant, though the bushes were deformed by the wind. There were scattered individuals of Erica scoparia subsp. azorica and Calluna vulgaris. 2. 'Laurelforest'
247
destroyed by m a n , but, according to Dansereau (1966), on steep escarpments on the northern side of the island extensive Laurel forests nearly 30 m high still survive. T h e 'relict Tertiary' flora is represented by Pittosporum coriaceum (endemic), Visnea mocanera (Canaries), Clethra arbórea (endemic), Sideroxylon marmulano (Canaries, Cape Verde), Heberdenia excelsa (Canaries), Picconia excelsa (Canaries), Persea indica (Azores, Canaries), Appollonias barbujana (Canaries), Ocotea foetens (Canaries) and Dracaena draco (Canaries, Cape Verde).
This type, which consists almost entirely of broad-leaved trees, appears to represent the climax up to c. 600 m . It is dominated by Lauras (Persea) azorica and Myrica faya, which reach a height of 6-7 m in favourable localities. Other species in the tree layer include Rhamnus The Canary Islands latifolia, Ilex perado subsp. azorica, Erica scoparia subsp. azorica, Viburnum tirtus, Vaccinium cylindraceum, Persea Refs.: Bôrgensen (1924); Bramwell (1976); Burchard (1929); Ceballos & Ortuño (1951); Ciferri (1962); Dansereau indica and Picconia azorica. T h e shrub layer is (1968); Follmann (1976); Kàmmer (1974, 1976); Künkel dominated by Myrsine africana. According to G u p p y the (1971, 1976); Lems (1960); Lindinger (1926); Schenck forests were originally m u c h taller than at present and (1907); Schmidt (1954, 1976); Sunding(1970, 1972, 1973a). Myrica faya, Laurus azorica and Erica scoparia subsp. azorica reached heights of 15 m , 15 m and 11 m The Canaries are a group of seven islands situated respectively. approximately 28° N . of the Equator (Fig. 25). Their total land area is 7273 k m 2 and the highest peak, on Tenerife, reaches 3718 m . Ecologically the islands can 3. 'Ericetum azoricae' be divided into two groups. In the first, the eastern F r o m 600 to 1500 m ; frost occurs above 760 m . Erica islands of Lanzarote and Fuerteventura, which he little scoparia subsp. azorica, which grows to a height of more than 100 k m from the African coast and d o not 4.5-6 m , is dominant, with Juniperus brevifolia subexceed an elevation of 650 m , have an arid climate. B y dominant. Juniperus, which has suffered greatly from contrast, the western islands (Gran Canaria, Tenerife, felling for its valuable timber, was probably formerly G o m e r a , Hierro, and L a Palma) are situated between dominant, and was considerably larger. G u p p y states 200 and 360 k m from the mainland and have a more that Taxus baccata formerly grew in the lower part of the oceanic climate. zone in considerable quantity but became extinct In general the Canaries have hot, dry summers and through excessive felling. w a r m , wet winters. Moisture is brought by the northeast trade winds, which are responsible for a cloud zone between 800 and 1500 m on the north side of all the 4. 'Callunetum' western islands. It is widely believed that fog precipitaF r o m 1500 m to the summit of Pico (2300 m ) ; between tion has a great influence on vegetation, but according 1500 and 1800 m Calluna is mixed with Erica scoparia to Karnmer the distribution of forest types on Tenerife is subsp. azorica. True Callunetum, which is an open not greatly influenced by this factor. T h e southern community on steep and often rather unstable slopes of sectors of the islands are in a rain shadow and are volcanic debris and little-weathered lava, is floristically mostly without a forest zone at m e d i u m altitudes. T h e very poor. T h e only c o m m o n species are Calluna extreme dryness of the eastern islands and the south of vulgaris, Daboecia azorica and Thymus caespititius. G r a n Canaria is partly due to the hot dry Sahara wind, the Harmattan, which sometimes blows for up to a week at a time. The eastern islands are too low to intercept the Madeira moisture-laden winds except at their highest points. T h e Réf.: co*ckereU (1928); Hansen (1969); Sjogren (1973, 1974, trade wind is responsible for differences of over 10°C 1978); Vahl (1905). between the north and south coasts of the larger islands. O n Tenerife above 1900 m snow lies for about five The Madeira islands he approximately 560 k m from the months of the year. African coast and 450 k m N . of the Canaries. Madeira, T h e flora of the Canaries amounts to c. 1800 species the main island, reaches an elevation of c. 2000 m . including m a n y introduced species. S o m e 460 species About 1140 species of flowering plants and ferns (25.5 per cent) are endemic, as are 17 genera. All 13 of have been recorded from the islands. O f these at least the Macaronesian genera occurring in more than one 250, and probably m a n y more, have been introduced. archipelago are found in the Canaries. About 120 species are believed to be endemic. T h e monotypic Chamaemeles is the only endemic genus. O f recent studies on the vegetation of the Canaries, Ceballos & Ortuño (1951) and Dansereau (1968) have Accounts of the vegetation are conflicting and difdescribed the Laurel forests of the western islands, and ficult to reconcile. M u c h of the vegetation has been
Vegetation ofthe floristic regions
248
!•• SLA GOMERA
o 0 Providence
. 1
Alphonse
COMOROS
J
\ jjjTromelin
•
/
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Juan de Noval
CD
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•
\
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Cargados Islands
/
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, ^ , Rodrigues
20' / Bassos d a 9 India
i > Europa
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* Mauritius
*s^°5
Reunion
/ /
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50°
60°
FIG. 27. Islands of the western Indian Ocean A single large star designates the, mainly Precambrian, Seychelles archipelago. Smaller stars represent the four volcanic islands of the C o m o r o group and the three volcanic islands of the Mascarenes. Solid circles indicate low islands, which are either raised coral-reef limestone islands or sand cays on sea-level reefs
20"
Other offshore islands
257
introduced and other species have escaped from cultivaThe earlier Floras of the Mascarene islands are out tion, so that the adventive flora (247 species) n o w of date and not always reliable, so that only very exceeds the indigenous. O n e family (Medusagynaceae) is approximate figures on endemism and floristics can endemic, as are the following ten genera: Valeria, be given. A n e w Flora (Bosser et al, 1976), however, Geopanax, Indokingia, Protarum, Deckenia, Lodoicea, is being prepared, and the vegetation of Réunion has Nephrosperma, Phoenicophorium, Roscheria and Verbeen recently described in considerable detail (Cadet, schaffeltia, the last six of which belong to Palmaceae. 1980). T h e original vegetation has been profoundly modified and over extensive areas totally destroyed. Mauritius Vesey-Fitzgerald and Jeffrey have reconstructed the Refs.: Brouard (1963); Sauer (1961, 1962); Vaughan (1968); original vegetation as follows: Vaughan & Wiehe (1937-47). 1. Coastal formations. In addition to mangrove and Ipomoea pes-caprae formations, the coast was fringed Mauritius is 62 k m long and 46 k m wide and has a total with coconuts (Cocos nucífera), with which were assoarea of 1865 k m 2 . The landrisesfrom the coastal plains ciated such typical sublittoral species as Scaevola to a rugged and precipitous central plateau (305-730 m ) sp., Cordia subcordata, Hibiscus tiliaceus, Hernandia on which are several extinct craters, and peaks which ovigera and Tournefortia argéntea. reach an altitude of 826 m . Vaughan & Wiehe recognise 2. Lowland rain forest. U p to 300 m on M a n é and two ecological zones, lowland and upland, which Silhouette. Canopy at about 30 m . Dominated by correspond to the coastal plains and the central plateau Imbricaría seychellarum and Calophyllum inophyllum. respectively. Associates: Dillenia ferruginea, Intsia bijuga, Valeria In the lowlands m e a n annual rainfall varies from seychellarum (Mahé only), Medusagyne oppositifolia 890 m m on the leeward side of the island to 1905 m m on (in deep clefts of granite masses, almost extinct). the south-east coast. In the uplands it ranges from 2540 3. Intermediate forest. F r o m 300 to 550 m on M a h é and to 4445 m m . Cyclones are frequent and devastate crops Silhouette. Canopy at about 18 m . Dominated by but do little damage to the indigenous forests, possibly Dillenia ferruginea and Northea seychellana. Asso- because of the efficient anchorage of the trees. ciates: Soulamea terminalioides, Colea seychellarum, Torrential rains which accompany cyclones, however, Campnosperma seychellarum, Riseleya griffithii, frequently cause landslides in the montane forests which Aphloia theiformis, Pandanus hornei. Most of the are rapidly colonized by exotic species. endemic species apparently occurred in this T h e flora of Mauritius is relatively small. Baker community. records 869 species for Mauritius and the Seychelles 4. Mossy montane forest. Above 550 m on M a h é . combined. Canopy at 12 m or lower. Dominated by Northea T h e indigenous vegetation has disappeared from seychellana. Associates: Roscheria melanochaetes, most of Mauritius. Even where it has not been destroyed Timonius seychellensis, Nepenthes pervillei. it is threatened by more vigorous, invasive, exotic species 5. Drier forest. In drier parts of M a h é , Silhouette, and such as Furcraea foetida, Ligustrum robustum, Ravenala Praslin. Dominant: Dillenia ferruginea. Prominent madagascariensis, Leucaena leucocephala (glauca), associates include Diospyros seychellarum, Dodonaea Albizia lebbeck and Psidium cattleianum, which prevent viscosa and Memecylon eleagni, and several endemic natural regeneration of the native species. palms including Lodoicea maldivica (Praslin and T h e lowland forests have been virtually destroyed. Curieuse only, the famous Coco-de-Mer), VerschafF r o m the accounts of early explorers it appears that feltia splendida and Deckenia nobilis. palm stands consisting of Latania lontaroides, Dictyosperma album and Hyophorbe sp. occurred where rainfall The lowland rain forest no longer exists and the other is less than 1000 m m per year. The moister forests were forest types survive only as small relict patches. Most of probably dominated by Diospyros tesselaria (endemic) the land is occupied by plantations or secondary and Elaeodendron orientale (also in Réunion and communities, of which the following are the most imporRodrigues) associated with Foetidia mauritiana, Stadtant: coppice of Cinnamomum zeylanicum, secondary mannia oppositifolia, Hornea mauritiana (endemic) and Albizia falcata forest, and thickets of the fern Dicranopteris linearis on worn-out, eroded, and denuded land. Terminalia bentzoe (Réunion, Rodrigues). T h e upland communities are somewhat better preserved. The following are the more important ones described by Vaughan & Weihe: 1. Swamp forest. Dominated by five endemic species of Pandanus. This group of three volcanic islands is situated on the 2. Sideroxylon thicket. T h e dominant, S. cinereum southernmost part of the Seychelles-Mauritius ridge (see (endemic), forms an 8-10 m high open canopy over a Fig. 27). T h e nearest large land mass is Madagascar, dense closed lower stratum of phanerophytes belong680 k m north-west of Réunion. The prevailing winds ing to ninety species. are the south-east trades which blow throughout the year, but more or less fitfully from December to April. 3. Upland forest. T h e main canopy at 18-21 (25) m is
The Mascarenes
258
Vegetation of the floristic regions
composed of Calophyllum eputamen, Canarium paniculatum {mauritianum), Mimusops maxima (endemic), M. petiolaris (endemic), Nuxia verticillata (also in Réunion), Sideroxylon cinereum a n d S. majus {Calvaría major, endemic). T h e boles are short and stout, about 1 m in diameter and branch at 10-15 m . There is a n intricate system of large roots o n the surface of the soil sometimes covering an area three or four times greater than the spreading canopy of the tree. T h e Sapotaceae have well-developed buttresses. 4. Mossy forest. This occurs o n M t Cocotte (744 m ) . Rainfall is m o r e than 4 0 0 0 m m per year. Clouds and mists are frequent. T h e 8-15 m canopy is very irregular, a n d is c o m p o s e d of Nuxia verticillata, Eugenia sp., Molinaea sp., Tambourissa sp., Aphloia theiformis, Turraea (Quivisia) oppositifolia etc. T h e trunks and branches are covered with a great variety of filmy ferns, mosses a n d hepatics. 5. Philippia thicket. T h e thicket, 4 m tall, occupies a small part of the plateau at 610-670 m , where it occurs o n little-altered lava. It is dominated b y Philippia abietina (endemic) associated with Phylica nitida {mauritiana, also o n Réunion) and Helichrysum yuccifolium (Réunion). Réunion
the dry side. These forests are remarkable for their low stature, with few trees exceeding 15 m in height. Characteristic species include Calophyllum tacamahaca, Grangeria borbónica and Pittosporum senacia, in addition to those mentioned under 2 above, which, in the moister forests, only occur at low altitudes. O n the windward slopes the forest diminishes in stature a n d at 1700 m is replaced b y 4 - 5 m high elfin thicket dominated b y Forgesia borbónica. 4. 'Tamarin' {Acacia heterophyllá) scrub forest. This type occurs in rain-shadow areas between 1200 and 2 0 0 0 m . T h e gnarled 8 - 1 0 m high Acacias are strongly dominant. Apart from a b a m b o o , Nastus borbonicus, other w o o d y plants are said to b e infrequent. Tamarin forest is less dense than the moister forests a n d is very susceptible to fire. It only regenerates o n bare soil, notably following fires. Although young plants are resistant to cyclones, the adults are easily blown over and u p to 80 per cent of a stand m a y b e destroyed in this w a y . 5. Ericoid communities. Between 2 0 0 0 and 2 5 0 0 m the vegetation of slopes is dominated b y Philippia montana, a n d above 2500 m b y Stoebe passerinoides. Associates include Hypericum revolutum {lanceolatum), Phylica nitida {leucocephala) and species of Psiadia and Philippia.
Refs.: Cadet (1980); Rivals (1952, 1968). Réunion, which is 75 k m long and 70 k m wide, is situated 780 k m E . of Madagascar and 200 k m S W . of Mauritius. T h e central massif culminates in Piton des Neiges at 3069 m . Rainfall at sea-level ranges from 425 m m per year o n the dry side of the island to 4290 m m per year on the south-east coast. O f the 630 species constituting theflorac. 480 are indigenous; 160 species belong to the Orchidaceae, but three-quarters of them are believed to be extinct. O f the remaining species, c. fifty are endemic to Réunion, but a m u c h larger number is shared with the other Mascarene islands. Vegetation zones are not clear-cut and some tree species have very wide ecological amplitudes. Thus, Aphloia theiformis and Nuxia verticillata range from sealevel to 2000 m and experience a m e a n annual rainfall from 800 to 7500 m m , and m e a n annual temperatures from 10° to 25° C . Agauria salicifolia grows from sealevel to 1000 m , above which it is replaced by A . buxifolia (also in Madagascar) up to 2500 m . Rivals recognizes the following plant communities: 1. Coastal. M a n g r o v e , halophytic, and Ipomoea pescaprae a n d Scaevola communities. 2. Dry, megathermic forest ( n o w destroyed). Below 400 m o n the dry side of the island. Important species were Elaeodendron orientale, Terminalia bentzoe {benzoin), Mimusops petiolaris, Diospyros melanida and Ocotea obtusata. 3. A complex of moister forests formerly ascending from sea-level to 1800 (2000) m o n the wet side of the island a n d occurring between 400 and 1200 m o n
Rodrigues Refs.: Balfour et al. (1879); Vaughan (1968); Weihe (1949). Rodrigues, the smallest (109 k m 2 ) of the Mascarene group, is situated 584 k m due east of Mauritius a n d rises to a height of 395 m above sea-level. T h e climate, which is relatively uniform, is not unlike that of the northwestern lowlands of Mauritius. M e a n annual rainfall o n the north-east coast is 1325 m m . O f the 145 indigenous species, 35 are endemic. T h e natural plant communities have disappeared and only individual plants remain. These are confined to the upper reaches of s o m e mountain streams or to sites where peculiar edaphic conditions have restricted cultivation, a n d alien vegetation has not yet gained a stranglehold. T h e original vegetation, apparently, w a s a low forest 10-15 m high with Sideroxylon galeatum {Calvaría galeota), Elaeodendron orientale, Mathurina penduliflora (endemic genus, Turneraceae), Diospyros diversifolia, Terminalia bentzoe, Foetidia rodriguesiana and the palm Dictyosperma album. P a l m stands, dominated b y Latania verschaffeltii and Hyophorbe verschaffeltii, a n d the drought-resisting screw-pine, Pandanus heterocarpus, seem to have been the characteristic vegetation of the coral plain o n the drier eastern coast. This plain is exposed to the southeast trade winds, which exert a profound influence o n the growth form of m a n y species. Shrubs—such as Carissa xylopicron—assume a cushion habit and rarely exceed a height of 40 c m .
Other offshore islands
Aldabra and other coral islands of the western Indian Ocean Refs.: Gwynne & Wood (1969); Renvoize (1975, 1979); Stoddart(1970). The scattered low islands of the West Indian Ocean are of two kinds: (a) raised reef-limestone islands best exemplified by the Aldabra group, which are mostly 3.5-8 m above m e a n low-tide level; (b) sand cays on sealevel reefs including the Farquhar group and African Banks. In recent years the flora and vegetation of Aldabra, the largest island (97 k m 2 ) , have been studied in great detail in connection with an investigation of the 150000 or so giant tortoises (Geochelone gigantea) which occur on the island. Comparative studies of some of the other islands have also been made. Although no comprehensive synthesis has yet appeared, this work has resulted in an extensive botanical literature, which is summarized below, together with floristic details, where known. Islands of the Mozambique Channel — Juan de Nova: Bosser (1952); Capuron (1966); Perrier de la Bâthie (1921¿). — Europa: Bosser (1952); Perrier de la Bâthie (19216). Amirantes group: 72 indigenous species; 25 introduced species. G w y n n e & W o o d (1969); Vesey-Fitzgerald (1942). — African Banks: Fosberg & Renvoize (in Stoddart, 1970); Stoddart & Poore (in Stoddart, 1970). — Remire: Fosberg & Renvoize, op. cit.; Stoddart & Poore, op. cit. — Desroches: Fosberg & Renvoize, op. cit.; Stoddard & Poore, op. cit. Aldabra group: c. 175 indigenous species of terrestrial vascular plants (c. 30 endemic); c. 85 introduced species. Fosberg & Renvoize (1980); Renvoize (1971); VeseyFitzgerald (1942); Wickens (1979). — Aldabra: Fosberg (1971); Hnatiuk & Merton (1979); Hnatiuk, Woodell & Bourn (1976); M a c N a e (1971); Merton, Bourn & Hnatiuk (1976); Stoddart & Wright (1967). — Assumption: Fosberg & Renvoize, op. cit.; Stoddart (1967). — Cosmoledo: Bayne et al (in Stoddart, 1970); Fosberg & Renvoize, op. cit.; Stoddart, op. cit. — Aslove: Bayne et al, op. cit.; Fosberg & Renvoize, op. cit.; Stoddart, op. cit. Farquhar group: 59 indigenous species. 13 introduced species.
259
— Farquhar: Fosberg & Renvoize, op. cit.; Stoddart & Poore, op. cit. — St Pierre: Stoddart (1967); Vesey-Fitzgerald (1942). — Providence: Stoddart (1967). Cargados Carajos: 48 k m 2 ; 17 indigenous species; 24 introduced species. Staub & G u e h o (1968). Isolated islands — Coetivy: 49 indigenous species; 16 introduced species. G w y n n e & W o o d (1969). — Gloriosa: 20 species. Battistini & Cremers (1972); Stoddart (1967). — Agalega: c. 60 species (J. Procter, pers. c o m m . in Renvoize, 1979). — Tromelin: 6 species. Staub (in Stoddart, 1970). O n m a n y of the islands the primary vegetation has been destroyed, either to m a k e w a y for coconut plantations, as on the Amirantes, or by guano-diggers, as on St Pierre and the islands of the Aldabra group excluding Aldabra itself. The vegetation on Aldabra, which has never been permanently settled, is little disturbed by m a n , although in places it has been profoundly modified by the resident tortoise population. T h e more important vegetation types on Aldabra are as follows: 1. Sclerodactylon macrostachyum tussock grassland and Sporobolus virginicus turf. Near the coast. 2. Mangrove. About 1-10 m or more high. Dominated by Avicennia marina, Bruguiera gymnorrhiza, Ceriops tagal and Rhizophora mucronata, with less abundant Sonneratia albida, Xylocarpus granatum and X. molucensis. 3. Pemphis acidula thicket. About 0.5-6 m high, usually on very rocky limestone. 4. Coconut {Cocos nucífera) groves. Introduced by m a n , locally naturalized. 5. Casuarina equisetifolia groves. Status uncertain. Possibly indigenous. 6. Mixed scrub. Widespread throughout the atoll, mostly 3-5 m tall, occasionally u p to 12 m . N o general dominants but the following are c o m m o n : Apodytes dimidiata, Canthium bibracteatum, Cassine aethiopica, Erythroxylum acranthum, Euphorbia pyrifolia, Ficus spp., Maytenus senegalensis, Ochna ciliata, Polysphaeria multiflora, Sideroxylon inerme, Terminalia boivinii. 7. Tortoise turf. W h e n intensely grazed 1-2 c m high, otherwise up to 15 c m . Composed principally of Dactyloctenium pilosum, Eragrostis decumbens, Panicum aldabrense, Sporobolus testudinum, and Bulbostylis basalis and other dwarf sedges.
Azonal vegetation xxii Mangrove, halophytic and fresh-water s w a m p vegetation
Mangrove
Mangrove
Herbaceous fresh-water swamp and aquatic vegetation
(mapping unit 77)
Halophytic vegetation
Refs.: Barbosa (1970, p. 121-5); Boughey (1957a, p. 679-80); Chapman (1977); Cole (1968, p. 72-5); Dale (1939, p. 7-8); Giglioli & Thornton (1965); Gledhill (1963); Gossweiler & Mendonça (1939, p. 70-2); Graham (1929); Hédin (1928); Hemming (1961, p. 64); G . Jackson (1964); Kassas (1957, p. 194-6); Kassas & Zahran (1965, p. 167-8); Keay (1953; 1959a, p. 11-13); Koechlin, Guillaumet & Morat (1974, p. 583-91); Letouzey (1968a, p. 239-40); Lugo & Snedaker (1974); Macnae (1963; 1968); Macnae & Kalk (1962a); Moll & Werger (1978); Naurois & Roux (1965); Pichi-Sermolli (1957); Pynaert (1933); Rabinowitz (1978); Rosevear (1947); Savory (1953); Trochain (1940); Walter (1971, p. 150-66); Walter & Steiner (1936); White (MS, 1975-76); White & Werger (1978). Phots.: Barbosa (1970: 14A.1); Boughey (1957a: 11-12); Gossweiler & Mendonça (1939: 4); Hemming (1961: 1); Karsten & Schenck (1915: 43); Kassas (1957: 1); Kassas & Zahran (1965: 4); Koechlin et al. (1974: 186-8); Macnae & Kalk (1962a: 1-4); Naurois & Roux (1965: 1-5); Pynaert (1933: 57-9); Walter (1971: 88-89, 98); Walter & Steiner (1936: 4-5, 8-11,13-14, 21-2). Profiles: Giglioli & Thornton (1965: 5); Gledhill (1963: 4); Naurois & Roux (1965: 5); Walter (1971: 87, 91); Walter & Steiner (1936: 7). Mangrove occurs only on shores where the vigour of the surf is broken by sand-bars or coral reefs or islands. It is most extensively developed o n the deltas of large rivers but also occurs in quite small bays and lagoons. Along rivers it can penetrate far inland. In West Africa, for example, it extends for 190 k m along the banks of the River G a m b i a . T h e seedlings of mangroves require quiet water on accrescent shores for their establishment. At the mouths of rivers the sea-water is diluted, sometimes for a considerable distance in a seaward direction, as in the case of largeriverslike the Niger. B y contrast the coastal mangroves growing away from the river inlets, in the shelter of coral reefs, are subjected to undiluted sea-water with an osmotic concentration of 24 atm. Mangrove attains its best development under a rainforest climate. In Africa the tallest stands, at the m o u t h of the Niger delta, are 45 m tall (Rosevear, 1947). O n s o m e shores, however, it extends far beyond the equatorial zone and even beyond the Tropics of Cancer and Capricorn, but it cannot tolerate frost. O n the west side of Africa mangrove is confined to the tropics. T h e
Mangrove, halophytic and fresh-water swamp vegetation
261
northernmost stand occurs near Tidra in Mauritania at cell sap. They can, however, in the absence of competi19°50'N., (Naurois & R o u x , 1965), and the southerntion, be grown on non-saline soil. Their leaves are succumost is near Benguela at 12°30'S. (Barbosa, 1970). A t lent and have water-storage tissue. T h e transpiration both places the m e a n annual rainfall is very low, c. rate is very low. Their roots are able to desalinate sea100 m m and 150 m m , respectively. O n the east side water to a high degree, but insufficiently so to prevent of Africa mangrove extends as far north as the Gulf some salt accumulation in their tissues. A m o n g African of A q a b a (30° N . ) and the Gulf of Suez (28° N . ) . genera only Avicennia is salt-excreting. In the dry The southernmost occurrences are near East London climate of East Africa the undersurface of the leaves of (33° S.). A. marina is densely covered with sodium chloride crystals. A t night the crystals absorb water hygroscopically T h e species occurring in West Africa are completely from the atmosphere and so dissolve. In more humid different from those in East Africa. There are five climates sodium chloride crystals are not formed. T h e species in West Africa, namely Rhizophora mangle, R. harrisonii, R. racemosa, Avicennia germinans (A. africana, salt is washed off by the rain. A. nítida) and Laguncularia racemosa. All five are also T h e three main environmental factors that influence widely distributed on the east coast of tropical America the occurrence and abundance of individual mangrove and neighbouring islands. species in relation to others are: The mangrove flora of East Africa is more diversi1. Frequency and duration of flooding with sea-water. fied, with nine species, namely Rhizophora mucronata, 2. Consistency of the soil; sandy or clayey. Avicennia marina, Sonneratia alba, Ceriops tagal, Bru3. T h e degree of admixture with fresh-water at river guiera gymnorrhiza, Xylocarpus granatum, X. molucmouths and the concentration of brackish water. censis, Lumnitzera racemosa and Heritiera littoralis. All Rainfall influences salt content, particularly that of these extend far towards the east and most reach the the rarely flooded higher regions. western Pacific Ocean. Mangroves frequently show a well-defined zonation of True mangrove species have either pneumatophores, species, but owing to the great variation from place to which are exposed at low tide, or are viviparous or place in the factors just mentioned and in floristic c o m almost so. M o s t African species show both these feaposition, and the wide climatic tolerance of most m a n tures. Species growing with mangroves which lack these grove species, there is n o generally applicable zonation. characteristics, e.g. Conocarpus erectus and Barringtonia racemosa, are referred to as 'mangrove associates'. Mangrove in West Africa In Rhizophora the stilt roots function as pneumatophores. Bruguiera, Ceriops and Lumnitzera have knee In the mangrove of the Niger delta, Rhizophora 'mangle' roots. In Xylocarpus and Heritiera the main roots are ( = R. racemosa"!, see below) covers 99 per cent of the laterally compressed and ribbon-like and their upper area and is said to attain a m a x i m u m height of 45 m and surface projects above the surface of the soil. In Avicena girth of 2.5 m above the 5 m long stilt roots (Rosevear, nia and Sonneratia erect pneumatophores arise from the 1947). The latter d o not penetrate the soil in the manner main subterranean roots. In Avicennia they resemble of normal roots, but, immediately below the surface of Asparagus shoots. T h e very large pneumatophores of the m u d , divide into innumerable rootlets of the thickSonneratia m a y be 75 c m high and 25 c m in diameter; ness of a piece of fine twine; and it is the vast developthey are used as floats for fishing nets. ment of these that completely alters the nature of the soil by displacement of the soft m u d . T h e tree therefore Rhizophora, Ceriops and Bruguiera are viviparous. stands upon a system of arches supported by a thick felt T h e embryo develops precociously and ruptures the raft of its o w n making and to which it is securely testa and pericarp, after which the hypocotyl undergoes anchored as it were by multitudinous tie-ropes. Accordenormous development. It contains the reserve food of ing to Rosevear this raft of rootlets, which is often a the seedling. In Rhizophora mucronata it reaches a length metre or more thick, spells the d o o m of the mangrove of almost one metre. W h e n the seedlings fall they m a y because Rhizophora can only reach its optimum size find anchorage beneath the parent plant, or m a y be diswhere it grows on newly deposited soft m u d . O n mature persed by ocean currents. Although not truly viviparous, mangrove sites the root systems of seedlings cannot the embryos of Avicennia, Laguncularia and Xylocarpus develop properly and nothing more than a low shrubby are well developed when they fall and emerge as seedtangle results. M a n y mangrove islands in the creeks conlings soon afterwards, from the fruit (in Avicennia and sist of an outer fringe of tall trees on freshly deposited Laguncularia, Gledhill, 1963; Rosevear, 1947) or seed m u d surrounding an inner core of this low tangle, which {Xylocarpus, F . White, pers. obs.). The indéhiscent fruits represents secondary or tertiary growth on the altered of Sonneratia are dispersed by bats (van der Pijl, 1957) soil. Rosevear states that the dominants of the low and and those of Lumnitzera and Heritiera by ocean curhigh mangrove are precisely the same tree. Savory (1953) rents. M o r e information on the dispersal and establishand Keay (1953, 1959a), however, recognize three ment of water-borne propagules of mangrove species has species of Rhizophora in the Niger delta and elsewhere been published by Rabinowitz (1978). on the coast of West Africa and eastern tropical A m e r According to Walter, mangroves are obligate haloica where they form separate communities within the phytes in that they accumulate sodium chloride in their
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Vegetation of the floristic regions
Rhizophora zone. T h e tall species, R. racemosa, is the commonest species and is the pioneer at the edge of the alluvial salt s w a m p to the exclusion of other species. R. harrisonii, which does not exceed a height of 6 m in Nigeria, is dominant in the middle areas, and the even smaller shrubby R. mangle is found only on the drier inner limit of the Rhizophora zone. Other workers have doubted the existence of as m a n y as three distinct biological species. Gledhill (1963) suspected a high degree of interfertility and hybridization between R. mangle and R. harrisonii. According to Breteler (1969), R. harrisonii is probably a hybrid between R. racemosa and R. mangle and sets little ripe fruit. J. B . Hall (in litt. 24 February 1977) shares this opinion and confirms that R. harrisonii scarcely forms any fruit in Ghana. In view of the uncertainty of field identifications the generic n a m e alone is used below. Behind the Rhizophora zone is the zone of Avicennia germinans with the grass Paspalum vagin*tum, which lies between the limits of the normal high tides and the spring tides, and is flooded twice monthly. Avicennia is normally a small tree which in the Niger delta rarely occurs within 24 k m of the sea. It can, however, in places grow in pure stands up to 30 m tall on the sea-board, as on Soden Island in the estuary of the Rio del Rey. This seems to happen when sand from the sea rather than silt from the river is deposited. Laguncularia bushes, which are almost totally submerged at high spring tides, appear to be the first colonizers in this situation and are subsequently suppressed by Avicennia. Avicennia can, like Rhizophora, occur in a stunted form, and the two are often found together in scrub mangrove, though it is exceedingly rare to find even a single specimen of this tree as a component of the main Rhizophora community of the open creeks. There is some evidence, however, that when Avicennia establishes itself as a pioneer on sandbanks as described above, it causes deposition of m u d which favours invasion by Rhizophora and hence the development of transitional mixed communities. The mangrove vegetation of Ghana, which occurs principally in lagoons, has been described by Boughey (1957a). West of Takorodi, where the rainfall is more than 1250 m m per year, all the lagoons are open, since they have a permanent outlet to the sea, and are thus inundated with sea-water by each tide. Most of the lagoons east of Takorodi, where the rainfall is lower, are closed, at least for the greater part of the year. They are open only for one or two months between June and September, the main rainy season, and not necessarily every year. Mangrove in open lagoons is dominated by Rhizophora, which is virtually confined to this habitat. Pandanus candelabrum also occurs and, in open places, the fern Acrostichum aureum. In closed lagoons at the high-water level reached during the seasonal flooding there is a fringe of scrubby Avicennia germinans associated with Laguncularia racemosa and Conocarpus erectus. Channels and pools of water which persist throughout the year in closed lagoons are fringed with Avicennia. T h e dried-out bed of the lagoon between the
channels is usually covered by a dense sward of Sesuvium portulacastrum mixed with Philoxerus vermicularis. T h e mangrove occurring on the Gambia River near Keneba 80 k m inland has been described by Giglioli and Thornton (1965). M e a n annual rainfall is 1125 m m and falls mostly between June and October. The inland swamps of the Gambia, unlike most littoral mangrove swamps, are not composed of regular belts of Rhizophora and Avicennia, backed by grasslands. The flatness of the land and the slow silting of the ancient Gambia flood plain have combined to form large areas of s w a m p , which ramify deeply into the surrounding country but nevertheless are reached by tidal water through a maze of meandering creeks. Each large or small watercourse is characteristically bordered, up to the limit of daily tidalflooding,by a gallery of Rhizophora, which in turn is usually surrounded to the mean limit of inundation by spring tides by open bushland of Avicennia germinans. Laguncularia racemosa is rarely seen. Rhizophora is usually the pioneer species. In some localities along the Gambia River, but not along its tributaries and associated swamps, Avicennia is the pioneer species. This happens only where sedimentation has been extremely rapid and a new high-shelving bank has been produced, which is barely covered by the daily tides. Such areas are usually immediately downstream of the mouth of a tributary. Behind or between the mangroves, there are isolated areas or continuous stretches of barren mud-flats which account for a quarter of the total area. They are 15 to 30 c m higher than the mangrove soil-level and are usually entirely without vegetation because they are desiccated during the dry season and their soils contain a high concentration of soluble salts, mainly chlorides and sulphates. Where the mud-flats are low enough to receive periodic flooding during the dry season from the high spring tides, they often, but not always, support perennial lawns of Sesuvium portulacastrum, and more rarely, at the end of the rains, seasonal lawns of Eleocharis spp. There are well-defined gradients in the depth of the water in the flood plains of the creeks, so that pure and highly zoned plant communities are produced. Rhizophora and Avicennia usually occur in remarkably pure stands. Mixing occurs only in the narrow ribbon-like ecotone between them, in which Rhizophora is being invaded and replaced by Avicennia. In West Africa Rhizophora reaches its northern limits just north of Saint-Louis in Senegal, and Avicennia at Tidra in Mauritania, where it forms an open bushland 2-2.5 m tall. South of Cameroun, mangrove is poorly developed, except at the mouth of the Zaire R . , where it shows the fullfloristiccomplement of West African species. Mangrove in East Africa All nine of the East African mangroves occur in Kenya, Tanzania, and Mozambique, but only Heritiera littoralis is confined to this part of the East African coast. T h e
Mangrove, halophytic and fresh-water swamp vegetation
other species, however, thin out rapidly to the north and south. Avicennia marina, Bruguiera gymnorrhiza and Rhizophora mucronata have the widest ranges and extend from the Red Sea to the Eastern Cape in South Africa. Ceriops tagal, Sonneratia alba and Xylocarpus granatum occur as far north as Somalia but reach their southern limits in Mozambique or the extreme north of Natal (Ceriops). Lumnitzera racemosa extends from Kenya to Natal. The zonation of mangroves in East Africa, which is m u c h more complicated than in West Africa, has been described for the Tanga region of Tanzania by Walter & Steiner (1936). They emphasize the fact that zonation shows m a n y irregularities and that the occurrence of the species is not directly related to distance from the outer and inner limits of the mangrove s w a m p but to the depth and salinity of the water and the texture of the alluvium. Sandbanks often occur at the outer limit of mangrove and here the water is very shallow. Elsewhere deep channels, which are only briefly uncovered by the receding tides, m a y extend a considerable distance inland. The ground level is not alwaysflatand a difference in micro-relief of only 20 c m , which is imperceptible to the eye, m a y be responsible for a change in dominance. For these reasons only a generalized zonation can be described. In the deepest water at the outer edge of the m a n grove Sonneratia alba is dominant and is so dense that it cannot be penetrated by a small boat. Inside the Sonneratia zone a Rhizophora mucronata zone is usually well developed. At river mouths Sonneratia, which is more closely associated with undiluted sea-water than other species, disappears almost completely from the outer zone and is replaced there by Rhizophora. Inside the Rhizophora zone there is usually a narrow zone of Ceriops tagal along small channels far into the innermost zone, the Avicennia marina zone. O n the sides of such channels, which are caused by erosion, zonation is telescoped into a few metres and Rhizophora, Ceriops, and Avicennia replace each other in rapid succession. In the wide innermost zone, which is not flooded daily with sea-water, but only at spring tides, Avicennia occurs almost pure as a small bush only a few metres tall. Towards the inner margin of the mangrove s w a m p Avicennia gets smaller and smaller until it occurs only as juvenile individuals in thick stands along scarcely perceptible channels with only scattered individuals on the intervening flats, which are only a few centimetres more elevated. Small h u m m o c k s in the Avicennia zone are colonized by Arthrocnemum indicum and Sporobolus virginicus. O n the landward side of the mangrove there is a vegetation-free sand-flat. This is related to the dry climate. T h e barren sand-flat is flooded only twice a year for a few days during the equinoctial spring tides. Subsequently the salt concentration increases considerably through evaporation. During the rainy season the salt is leached out. It appears that neither halophytes nor glycophytes can grow under these conditions. At the
263
lower limit of the barren zone, Suaeda monoica is often frequent, or there m a y be scattered bushes of Lumnitzera racemosa. In the more open parts of the Avicennia fringe, Suaeda monoica is often associated with Arthrocnemum indicum, Sesuvium portulacastrum and the grasses Sporobolus virginicus, Paspalum vagin*tum and Dactyloctenium geminatum. In the Tanga region Bruguiera gymnorrhiza does not form a distinct zone but occurs scattered with Rhizophora and Ceriops. It also penetrates a considerable distance inland along rivers. The fern Acrostichum aureum is often abundant near river mouths but only in the inner zones where the water is brackish. It is completely absent from mangrove bathed with pure sea-water. In the innermost mangrove zone Avicennia is always bushy. It can also occur as a large tree at the outer edge of mangrove, though it never forms a continuous zone there. It seems that in this situation it always occurs on sandy alluvium. Mangrove species also grow on coral rock but because of the extreme irregularity of the surface they show n o zonation there. In Kenya the behaviour of the mangrove species is similar to that just described for Tanga in Tanzania, though their relative abundance varies greatly according to local conditions. Rhizophora mucronata is the most abundant species, covering 70 per cent of the area. It often occurs pure or with a few scattered Bruguiera. The most extensive and luxuriant mangrove s w a m p occurs in the L a m u archipelago at the mouth of the Tana River. Further south in Kenya in Gazi Creek at the mouth of the very m u c h smaller Kidogoweni River, the alluvium is mostly sandy, and Rhizophora is largely confined to the m u d d y banks of the main channels, which receive a copious supply of fresh-water; Xylocarpus granatum is locally abundant and there are scattered individuals of Bruguiera. Smaller channels which have sandy banks and receive less fresh-water are fringed by a narrow zone of Ceriops and Lumnitzera with scattered Avicennia up to 11 m tall, and Heritiera. T h e extensive sandy flats between the smaller channels support open Avicennia bushland 2-5 m in height. In Eritrea, mangrove dominated by Avicennia occurs on sandy m u d overlying coral rock in shallow bays which are partly land-locked and sheltered from the full force of the tides by coral reefs (Hemming, 1961). In the Sudan Avicennia marina is the principal species. Where the water is shallow and the substrate compact, camels browse the leaves and shoots and the vegetation is noticeably thinned. In deeper water Avicennia forms dense thickets. In the extreme south of the Sudan, Avicennia is associated with Rhizophora mucronata and Bruguiera gymnorrhiza. Nearly all the East African mangrove species occur at the mouth of the Zambezi delta in Mozambique, but mangrove s w a m p is relatively restricted there and extends only 15 k m along the main channel, although salt-water goes up m u c h further (Macnae, 1968).
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Vegetation of the floristic regions
O n Inhaca Island at the extreme southern end of Mozambique, mangrove occurs on the sheltered shores. Zonation occurs along creeks but not in long-shore m a n grove. O n open slopes and at the mouths of estuaries Avicennia marina is the principal species probably because of the sandy soil. Upstream it is replaced by Rhizophora mucronata. The greater part of the mangrove s w a m p is m a d e up of Ceriops tagal in the drier parts, and Bruguiera gymnorrhiza where the water-table is at or near the surface. O n the landward fringe of the Ceriops zone, Lumnitzera racemosa is frequent and Xylocarpus granatum occurs rarely. Avicennia reappears locally in the form of low stunted bushes at the landward edge of the s w a m p ; elsewhere there m a y be bareflatswith saline efflorescence (Macnae & Kalk, 1962a). O n all Inhaca shores, Avicennia marina is the principal colonist. It grows on sandy beaches and on m u d d y ones in which drainage is improved by the inclusion of shell debris. O n c e established the pneumatophores cause the accumulation of silt around them, followed by the invasion of Ceriops and Bruguiera, the seedlings of which require shade for their establishment. Further accumulation of m u d and the formation of mangrove peat causes waterlogging of the soil. Avicennia dies back under these conditions. Rhizophora on the other hand will only grow into a tree in waterlogged soil, but only if the salinity is below that of normal sea-water (Macnae & Kalk, 1962a). In South Africa, mangrove is not extensively developed but has a scattered distribution along the east coast. It is absent from the mouths ofriverswhich run to the sea through deep gorges and have only a narrow flood plain. Mangrove is also absent from the mouths of rivers which become cut off from the sea by sand-bars during the dry season. Such estuaries are dominated by Barringtonia racemosa and Hibiscus tiliaceus (Macnae, 1963). Five species, Avicennia marina, Bruguiera gymnorrhiza, Rhizophora mucronata, Ceriops tagal and Lumnitzera racemosa, occur at Kosi Bay in Zululand. Ceriops and Lumnitzera reach their southern limits there. T h e behaviour of Lumnitzera in unusual. Elsewhere it occurs at the landward margin of mangrove swamps and its roots are submerged only at the highest spring tides. At Kosi Bay it occurs with Ceriops and Bruguiera in all thickets and is to be found on the seaward fringe as well as on the landward. Its occurrence at low intertidal levels m a y be explained by the absence of m u d and the good drainage provided by a sandy substrate. The same factor m a y account for the abundance of Avicennia in Natal and the rarity of Rhizophora. Avicennia is normally the pioneer species in southern Africa and is capable of colonizing stable sandy shores. Bruguiera is the species most tolerant of fresh-water and reaches its most luxuriant development in areas of low salinity. Rhizophora is more dependent on m u d and only occurs abundantly in those estuaries where mud-banks are well consolidated and border creeks or channels.
Avicennia, Bruguiera and Rhizophora extend as far south as Jhe mouth of the Kei River In Madagascar mangrove covers 217600 hectares (Koechlin et al., 1974). Nearly all of this occurs on the western coast. Mangrove is extremely localized on the exposed eastern coast, where it is confined to a few sheltered estuaries. All of the nine species of mangrove which are found in East Africa also occur in M a d a gascar, where their ecology appears to be similar. Mangrove is of considerable economic importance. It provides poles and planks for house-building and small planks for boat-building. In East Africa poles of Heritiera littoralis are m u c h used for the masts of Arab dhows. Mangrove is also an important source of fuel. In Nigeria the coastal towns were formerly dependent on it. In East Africa large quantities of Rhizophora mucronata bark were formerly exported for tannin, while the local people still use mangrove tannin for preserving their fishing nets, ropes and sails. Mangrove swamps in West Africa also provide breeding-places for the salt-water vector of malaria, Anopheles gambiae melas, but after appropriate reclamation measures they can be used for growingriceand other crops.
Herbaceous fresh-water s w a m p and aquatic vegetation (mapping units 64 and 75) Refs.: Boughey (1963a); Eggeiing (.1935); Germain (1965, p. 217-44); Greenway (1973); Howard-Williams & Walker (1974); Léonard (1952; 19696); Lind (1956è); Lind & Morrison (1974, p. 102-27); Lind & Visser (1962); Mitchell (1978); Seagrief (1962); Thompson (1976); Van der Ben (1959); Van Meel (1952, 1953, 1966); Wild (1961); Wild & Barbosa (1968, p. 64). Phots.: Eggeiing (1935: 1-4); Germain (1965: 1-8); Lind (19566: 1, 2); Lind & Morrison (1974: 30-5); Seagrief (1962: 1, 2); Thompson (1976: 28); Van der Ben (1959: 1-10); Van Meel (1952: 14, 15, 18; 1953: 1-13). Profiles: Lind & Morrison (1974: 3, 4); Thompson (1976: 26). Throughout the wetter parts of tropical and subtropical Africa water accumulates in depressions, where it gives rise to s w a m p s and lakes. In the Guineo-Congolian Region most s w a m p y areas are covered with s w a m p forest (page 82). B y contrast, reed-swamp and aquatic communities are relatively restricted. Outside the Guineo-Congolian Region most of the shallower lakes, except those which are strongly saline, have a wide belt of reed-swamp, of which the main constituent is Cyperus papyrus or 'papyrus', the tallest m e m ber of the Cyperaceae. R e e d - s w a m p also occurs less extensively in sheltered bays on the shores of deeper lakes and in the backwaters and lagoons of the largerrivers.Conditions favourable for the development of reed-swamp are widespread in East and South tropical Africa, where they have developed following warping of the earth's crust and other tectonic movements, in places associated with rift formation.
Mangrove, halophytic and fresh-water swamp vegetation
265
In Uganda, s w a m p occupies 6 per cent of the total The grass Vossia cuspidata is the most characteristic land surface, of which the largest areas are produced by pioneer of reed-swamp and is usually abundant at the the Nile, especially where the Victoria Nile flows outer edge of the papyrus zone. Its stems, which are up through Lake Kioga. Further north, in the Sudan, the to 6 m long, he on the water, and with other prostrate White Nile with its tributary the Bar el Ghazal forms the species such as Ludwigia and the fern Thelypteris striata largest s w a m p in the world, the Sudd, which covers an form strongfloatingmats which will bear the weight of a area of 150000 k m 2 and extends for more than 600 k m m a n . The stout rhizomes of Cyperus papyrus push out from north to south and a similar distance from east into this mat and so extend the area of reed-swamp. to west. Other species rooted in the Vossia mat include the grasses Echinochloa pyramidalis, E. scabra (stagnina), T h e largest s w a m p areas in West Africa are on the Oryza longistaminata (perennis) and Paspalidium gemishores of Lake Chad and in the valley of the Upper natum, and the sedges, Eleocharis acutangula and Scirpus Niger south of Tombouctou. inclinatus. There are also m a n y swamps in the Zambezian Region, principally the Okavango, Busanga, and In deeper water beyond the reed-swamp there are Lukanga swamps, and those associated with Lakes normally two communities, the submerged community U p e m b a , M w e r u , M w e r u Wantipa, Bangweulu, Shirwa and thefloating-leafcommunity. and Chiuta. Smaller swamps fringe the flood plains of The most characteristic members of the submerged the Zambezi and Kafue Rivers. community are Ceratophyllum demersum, Hydrilla verticillata, Lagarosiphon spp., Najas spp., Ottelia ulvifolia, According to Debenham (1952) the vegetation growPotamogetón schweinfurthii, Utricularia spp. including U. ing in a s w a m p is not merely a concomitant of the water foliosa, Vallisneria aethiopica, V. spiralis and the charoconditions but sometimes is the cause of them. In some phyte Nitella. They show a clearly marked zonation in swamps there was no real ponding by physical features relation to water depth. Ceratophyllum, for instance, until the growth of the vegetation itself caused sufficient grows in water up to 8 m deep, while Potamogetón obstruction to hold back the water. schweinfurthii rarely tolerates more than 3 m . The dominants of reed-swamp are normally rooted There are two types of plants withfloatingleaves, in the soil beneath the water, but some, especially namely those which are rooted in the m u d and those papyrus and the grass Vossia cuspidata, also extend outwhich are free-floating. A m o n g the former are species of wards as afloatingmat at the edge of the s w a m p , which Nymphaea, including N. caerulea and N. lotus, Potamofrequently becomes detached and formsfloatingislands. getón richardii and Nymphoides indica. The principal In deeper water beyond the reed-swamp, communities of free-floating species are Azolla africana, Lemna spp. submerged and free-floating aquatics occur. including perpusilla, Trapa natans, Wolffia arrhiza, EichCyperus papyrus, the commonest dominant of reedhornia crassipes, Pistia stratiotes and Salvinia molesta. s w a m p , is widely distributed in tropical and south Africa These species are readily dispersed by water-movements and in Madagascar. It does not occur above 2300 m . and by wind. W h e n they are transported to favourable Formerly there were extensive growths in the Egyptian conditions they increase explosively, and all seven can Nile, where for centuries in dynastic time it provided the become troublesome, especially Eichhornia, Pistia and basis of a paper industry and was widely used for boatSalvinia. making, cordage, matting, food, and medicine. It has long been thought to be extinct there, but has been Eichhornia crassipes, the Water Hyacinth, is indigenrediscovered (El Hadidi, 1971). ous to tropical America but was introduced to Africa about 100 years ago. It is n o w widespread and is a Cyperus papyrus is a vigorous species which usually serious pest in m a n y places since it interferes with naviforms almost pure stands. It is very fast-growing and can gation in largerivers,blocks irrigation canals, and fills attain its m a x i m u m size of 5 m in 10 weeks. Other reeddams. Because of its rapid vegetative reproduction it is s w a m p dominants are m u c h more localized. In East Africa, Phragmites australis and P. mauritianus are com- difficult to eradicate, and expensive control measures in Zaire and the Sudan have been unsuccessful. A n indimonest in silted areas and in lakes of volcanic origin. Typha australis, T. latifolia and Cladium mariscus locally genous species, E. natans, which is distributed from Senegal to Angola and Zimbabwe, is not normally replace papyrus at higher altitudes. In some places in troublesome but is regarded as a potential pest in the East and South tropical Africa belts of the grasses LouGambia. detia phragmitoides and Miscanthus violaceus occur in shallow water on the landward side of papyrus Pistia stratiotes, the Water Lettuce or Nile Cabbage, swamp. has a pan-tropical distribution and has been k n o w n in Egypt since the time of Pliny ( A D . 77). It is widespread The principal associates of papyrus are Cyperus hasin tropical Africa but is not so dangerous as Eichhornia. pan, Dissous rotundifolia, Hibiscus diversifolius, Impatiens irvingii, Ipomoea spp., Ludwigia erecta, L. leptocarpa, It is said to be troublesome in the Gambia, the Niger L. octovalvis, L. stolonifera, Limnophyton obtusifolium,Delta, the Upper Nile, parts of Kenya, and the littoral regions of Angola and Mozambique. Melanthera scandens, Melastomastrum segregatum, Salvinia molesta, a free-floating water-fern, has been Mikania cordata, Polygonum pulchrum, P. salicifolium, confused with S. auriculata. The former is n o w thought P. strigosum, Thelypteris striata and Vigna luteola.
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Vegetation of the floristic regions
to be a sterile triploid hybrid which probably originated in cultivation and m a y have been introduced to Africa as an aquarium plant. Little was k n o w n about it before the construction of the Kariba d a m in the Zambezi Valley in 1959, following which it rapidly became an extremely serious weed on Lake Kariba where it covered large areas with a blanket 25 c m thick. T h e latter provided a substrate for sudd-forming species such as Vossia cuspidata and Scirpus cubensis. Sahinia has subsequently declined somewhat but is still a troublesome pest. Its explosive development is described by Boughey (1963a) and Mitchell (1978). In Uganda Miscanthus violaceus occurs in the inner part of the papyrus zone but it also forms a distinct zone in shallower water from which papyrus is absent. In the inner part of the Miscanthus zone another tall tussock grass, Loudetia phragmitoides, is co-dominant. W e t hollows alternate with the grass tussocks. The associates of Miscanthus are more diversified than those of papyrus and include Cyperus haspan, Dissous incana (canescens), Fuirena umbellata, Hypericum lalandii, Leersia hexandra, Paspalum scrobiculatum, Polygonum spp., Scleria nyasensis, S. nutans, Smithia elliotii, Thelypteris confluens, Tristemma incompletum and Utricularia gibba. Towards the landward margin of reed-swamp a narrow zone of shrubs and small trees adapted to s w a m p conditions often occurs. T h e principal species are Aeschynomene elaphroxylon, A . pfundii, Kotschya africana, Mimosa pigra and Sesbania sesban, often with scattered juveniles of swamp-forest trees such as Syzygium cordatum, Ficus verruculosa and Ficus congensis. R e e d - s w a m p fringing larger lakes, however, is not normally replaced by s w a m p forest, probably because the succession is arrested by the great changes in waterlevel which periodically occur. Sphagnum is normally absent from papyrus swamps, but is often found in Miscanthus swamps, especially those at higher altitudes. In the Zaire basin permanent s w a m p along all the major rivers is almost exclusively dominated by Vossia cuspidata. Its principal associates, except for Polygonum acuminatum, are grasses, namely Brachiaria mutica, Panicum subalbidum, Echinochloa pyramidalis, Leersia hexandra, Echinochloa scabra and Panicum parvifolium. Most openings in the s w a m p forest are dominated by Cyrtosperma senegalense, a giant aroid, which largely replaces Cyperus papyrus in the Zaire basin. There have been several plans for the draining and cultivation of African reed-swamp or the harvesting of their natural growth for manufacturing paper or hardboard. The dangers involved in exploiting such a fragile ecosystem are stressed by T h o m p s o n (1976).
Halophytic vegetation (mapping unit 76) Refs.: Emberger (1939, p. 147-8); Giess (1971, p. 9); Greenway (1973, p. 57); Lind & Morrison (1974, p. 174-5); Seagrief & Drummond (1957, p. 110-11); Symoens (1953); VeseyFitzGerald (1955a; 1963, p. 261-3; 1970); Walter (1971); Wild & Barbosa (1967, p. 61-2, 67-8). Phots.: Giess (1971: 46-9); Vesey-FitzGerald (1963: 4, 7). Saline soils are frequently found in arid and semi-arid regions where rainfall is insufficient to transport salts formed, during weathering, to the sea. Their distribution, however, is also partly determined by geology, in that locally they occur in wet regions around springs which bring soluble salts to the surface from salt-containing strata. A n intermediate condition occurs extensively under a m e a n annual rainfall of 250-1000 m m in parts of East Africa where the salts which are deposited in lake basins andrivervalleys are derived from volcanic depositsrichin sodium. Only a relatively small number of plant species, k n o w n as halophytes, grow on saline soils. T h e most typical halophytes absorb soluble salts, especially sodiu m chloride, which reaches high concentrations in the cell sap of the leaves. Non-halophytes cannot tolerate high internal concentrations of sodium and die. A comprehensive description of the vegetation of coastal saline soils that are influenced by sea-water is beyond the scope of this book. S o m e types, however, are briefly mentioned in the section on mangrove. T h e halophytic vegetation of the Sahara desert is dealt with in Chapter XVII and that of the KarooN a m i b Region in Chapter V I . In parts of the Karoo, especially the 'vloere', the vegetation consists essentially of halophytes, but halophytes are also widely distributed elsewhere in the Karoo. Indeed, species of the genus Mesembryanthemum sensu lato, which occur throughout the Karoo, always have high osmotic values and accumulate chloride even on non-saline soils. The vegetation on saline soils in the Somalia-Masai Region is briefly mentioned in Chapter IV. T h e vegetation on inland saline soils is physiognomically varied and includes grassland, wooded grassland, shrubland, and bushland. In the Maghreb halophytic vegetation occurs chiefly in the arid and semi-arid 'étages'. The principal species are Atriplex halimus, Lycium intricatum, Suaeda fruticosa, Salsola longifolia (oppositifolia), S. vermiculata, Asparagus stipularis, Anabasis aphylla, Peganum harmala, Artemisia herba-alba, Asphodelus fistulosus and species of Frankenia, Mesembryanthemum, Sphenopus, Lepturns and Ai'zoon. Non-halophilous therophytes are also plentiful. In more permanently moist depressions the following are frequent: Tamarix spp., Juncus maritimus, J. acutus, Statice spp., Scirpus holoschoenus, Spergularia marítima and Plantago coronopus (see also page 230). O n the eastern side of tropical Africa halophytic vegetation occurs in most of the lake basins in the Eastern Rift, principally Lakes Turkana, Hannington,
Mangrove, halophytic and fresh-water swamp vegetation
Nakuru, Elementeita, Magadi, Natron, Manyara, Eyasi and R u k w a . T w o lakes, however, L . Baringo and L . Naivasha, are m u c h less saline and probably o w e their freshness to subterranean outlets. Lake M w e r u Wantipa in north-east Zambia also lies in a d o w n faulted depression with internal drainage, which supports halophytic vegetation. S o m e of the lakes m e n tioned above are surrounded by extensive salt deposits that are the basis of an important industry. T h e chief plants around the saline lakes in K e n y a and U g a n d a are Cyperus laevigatus, Sporobolus spicatus and Dactyloctenium sp. T h e halophytic vegetation in the Lake R u k w a basin in Tanzania is chiefly grassland (Vesey-FitzGerald, 1963). It covers extensive level plains which are liable to be inundated by the lake during periods of high water, but which themselves extend over the dry bed of the lake w h e n the latter dries up. T h e soil is highly alkaline ( p H 8.0-9.6) and the surface is impregnated with soda. T h e grassland can be divided into the following three zones: 1. Beach zone. T h e fringe of the lake at its m a x i m u m extent is marked by a pure stand of 1-2 m tall tussocks of Sporobolus robustus. It never grows in water. Under dry conditions it persists for an indefinite period, making a clearly defined but obsolete beach zone which is often remote from the contemporary perimeter of the water in the lake. 2. Alkaline swamp. Because the beds of alkaline lakes are very flat, extensive areas are shallowly flooded w h e n the basin is full. This alkaline s w a m p is colonized by a single species, Diplachne fusca, which forms a dense and even mat up to 2 m in height and extends for m a n y kilometres. If the s w a m p dries up, even for a period of years, as often happens, the D . fusca mat remains in occupation of the ground for an indefinite period, but only grows to a height of 50 c m . W h e n growing in water D . fusca remains green throughout the year but when the lake has receded it dries u p and becomes burnt during the dry season. If m a n - m a d e fires do not consume the dry mat, lightning frequently ignites it at the onset of the rains, but even if burnt every year the Diplachne mat still persists. 3. Alkaline flats. The lake bed itself (as opposed to the lake swamps just described) shows a cyclical succession. The two grass species that are principally involved replace each other as the bed is successively flooded or dries out. W h e n the lake dries out a crust of saline soil is left on the surface but moist m u d remains below the surface mulch. This newly exposed surface is colonized by Sporobolus spicatus, which spreads rapidly by runners and produces a perennial sward. T h e latter persists during the most prolonged dry periods and is not
267
replaced by any other species. Another grass of similar growth-form, Psilolemma (Odyssea) jaegeri, m a y occur with S. spicatus but its tufts do not spread when the ground is dry. W h e n , however, the lake is rising this species colonizes vast areas of flats that are shallowly flooded with w a r m water in which the soda crust is dissolved. Under these conditions the S. spicatus m a t rots away and Psilolemma replaces it. In the W e m b e r e depression south of Lake Eyasi in the East African Rift Valley, alkaline s w a m p is dominated by Diplachne fusca grassland. Scattered Tamarix and 'salt bushes' (Chenopodiaceae) occur along the edges of small drainage lines. M a n y of the flat valleys in the drier parts of Tanzania have alkaline soils. This is particularly true of the Pangani River, the headwaters of which rise from the volcanic deposits of M t s M e r u and Kilimanjaro, which weather rapidly and release large amounts of salt into the drainage water. Saline soils on the flood plain are dominated by grasses and Sesbania sesban and the tree Acacia xanthophloea (but see page 30). Prominent halophytes include Salvadora pérsica, Sporobolus robustus, Suaeda monoica and Triplocephalum holstii. In South tropical Africa saline soils are more localized than in East Africa. T h e principal occurrences are in the Etosha depression in Namibia, the Makarikari basin in Botswana, and in the Changane valley in Mozambique. T h e Etosha pan itself is quite barren but it has a fringe of halophytic vegetation consisting principally of Suaeda articulata, A triplex vestita, Sporobolus spicatus, S. tenellus, S. virginicus and Odyssea paucinervis, surrounded by a dwarf-shrub zone characterized by Acacia nebrownii, Monechma tonsum, M . genistifolium, Leucosphaera bainesii, Petalidium engleranum and Salsola tuberculata. T h e Makarikari pan is surrounded by a narrow fringe of grassland dominated by Sporobolus spicatus and Odyssea paucinervis. Halophytic communities are widespread in the valley of the Changane, a tributary of the Limpopo. Rainfall amounts to 400-600 m m per year. In moderately saline zones there are grasslands of Acacia nilotica subsp. kraussiana. W h e n the salinity increases, the grasses Eriochloa meyerana, Sporobolus nitens and Aristida adscensionis form discontinuous patches with extensive bare areas in between. Near theriveritself salinity is high and species of Arthrocnemum, Salicornia, Atriplex and Suaeda predominate. Symoens (1953) has described the M w a s h y a salt-flats in the Lufira Valley 30 k m S W . of Lukafu in Upper Shaba. They are dominated by Juncus maritimus and Sporobolus cf. virginicus, two coastal species which have a very sporadic distribution in the interior of Africa.
Glossary and index of vernacular names of vegetation types and habitat
Note. Definitions, where given, are generally brief, since the meaning of most terms should be apparent from the text (to which reference is made).
classified separately. Elfin thicket does not comfortably fit any of the categories of conventional classifications of vegetation. Hence its continued use seems appropriate, notwithstanding its somewhat whimsical connotation. E R G , 218, 220. A sand desert usually in the form of dunes.
A L F A , 216, 229ff. Esparto grass, Stipa tenacissima. B A N , 116. A treeless plain in Somalia. B A R C H A N , 220. A crescent-shaped mobile sand dune devoid of vegetation. BATEKE. See TEKE
B A T H A , 159. A type of spiny dwarf shrubland in the Mediterranean Region. B O W A L (pi. B O W E ) , 108. Ironstone outcrop covered with open, seasonal marsh vegetation. B R O K E N V E L D . Applied by South African botanists to a landscape dominated by dwarf shrubs, grasses, and succulents with scattered larger bushes and occasional small trees, usually occurring on flattish, gravelly plains or rugged, rocky mountains. Walter (1971, p. 391), however, defines it as grassland interrupted by areas with a different vegetation. B R O U S S E TIGRÉE, 23, 27, 213. A pattern of arcs or stripes of vegetation alternating with bare areas in arid and semi-arid regions. CHIPYA, 60, 96. Fiercely burning wooded grassland on Central African Plateau (from Bemba 'cipya').
F A D A M A , 107. A grassy flood plain in West Africa. FiRKi, 108. Seasonally flooded wooded grassland in the Chad basin. F Y N B O S , 32, 41, 48, 49, 132ff, 159. The characteristic sclerophyllous vegetation of the Cape Region. Most fynbos is shrubland, less often bushland or thicket. GARRIC See GARRIGUE
G A R R I G U E , 49, 159. A type of short Mediterranean shrubland. G U E L T A , 222. A pool, more or less temporary, in the bed of a wadi, usually sheltered in a canyon. H A L F A . See A L F A
H A M A D A , 218, 220. A stone desert incised by dry valleys (wadis). H I G H V E L D , 51, 64, 194. A n Afrikaans word applied to the plateaux surfaces of the interior of southern Africa. In this work restricted to the eastern part of the transition zone between the Zambezian and Karoo-Namib Regional Centres of Endemism. ITIGI THICKET, 48, 97.
CHOTT. See SHATT
C I T E M E N E , 92. A type of slash-and-burn shifting cultivation practised mainly in miombo woodland on the Central African Plateau. C R A M - C R A M , 216. Cenchrus biflorus. D A M B O , 61, 99ff. A seasonally waterlogged depression on the Central African Plateau covered with grassland. ELFIN T H I C K E T , 83, 168. The term 'elfin woodland' was introduced by the translators of Schimper's Pflanzengeographie (Schimper, 1898, 1903) as the English equivalent of 'Krummholz'. Tropical examples of the latter were described by Schimper as montane woody communities dominated by dwarf trees with short, thick, gnarled trunks which often are almost horizontal or, at least, heavily leaning. In English, the term has subsequently been applied to various types of stunted, montane, cloud 'forest' with abundant epiphytes, especially liverworts and mosses. African examples, however, fit the definition of thicket, rather than woodland, as used in the present work. In Africa, elfin thicket occurs on the summits of rather low mountains at lower altitudes than the general upper limit of the forest belt. Ericaceous bushland and thicket are
K A R A L . See FIRKI
K A R O O , 137ff. A Hottentot word meaning bald, applied to the essentially treeless semi-desert parts of South Africa which are mainly dominated by succulent communities and dwarf malacophyllous shrublands. MABWATi, 172. A type of dry evergreen forest in Zaire and Angola. MACCHIA. See MAQUIS
M A Q U I S , 41, 49, 159. A type of Mediterranean sclerophyllous shrubland, usually tall, often impenetrable. The term is frequently misapplied to sclerophyllous shrublands (e.g. fynbos, q.v.) in other parts of the world. MATESHi, 96. A type of Zambezian dry evergreen thicket. MATTORAL, 149. All woody non-forest vegetation in the Mediterranean Region. M A V U N D A , 90. Zambezian dry evergreen forest dominated by Cryptosepalum pseudotaxus. M B U G A , 116. Water-receiving depressions in East Africa covered with grassland and acacia-wooded grassland on seasonally saturated, black, cracking clays. Mostly occurring at lower altitides and under a drier and hotter climate than dambos.
270
Glossary and index of vernacular names of vegetation types and habitat
MIKWATI, 173. A type of (chiefly secondary) wooded grassland in Zaire. MiOMBO, 54, 57, 60, 61, 92, 181. A kind of woodland in the Zambezian Region dominated by species of Brachystegia and related genera. M O P A N E , 54, 61, 62, 93, 143, 191. A vernacular name applied to Colophospermum mopane and vegetation dominated by it. M U H U L U , 91. A kind of Zambezian, dry evergreen forest. M U T E M W A , 90. The shrub layer of dry deciduous forest dominated by Baikiaea plurijuga. It usually forms a 5-8 m high thicket. N O O R S V E L D , 141. Vegetation in the Eastern Cape intermediate between bushland and shrubland, and dominated by Euphorbia coerulescens (Noors). P A P Y R U S , 264. The giant sedge Cyperus papyrus and vegetation dominated by it. PARAMO, 169. PENGBELE. See BOWAL PSEUDOSTEPPE, 173.
Q O Z , 102. A consolidated sand dune. REG, 218, 220, 221. A gravel desert. R E N O S T E R V E L D . See R H E N O S T E R B O S V E L D
R H E N O S T E R B O S V E L D , 132, 135. Cape shrubland dominated by Rhenosterbos, Elytropappus rhinocerotis. S A B K H A (also transliterated S E B K H A or S E B K R A ) , 222. The class-
ical Arabic term for saline or swampy land. It is used for the salt lakes of inland drainage basins, which are periodically filled but usually dry. 'Sebkha' is the form most often used in botanical literature. See also S H A T T . S A H E L . The original meaning in Arabic is seashore, but it was subsequently extended to gravel plains and to the northern margins of the Sahara (P. J. Stewart, pers. c o m m . ) . It was first applied in its phytogeographical sense by Chevalier (1900) to the southern fringes of the Sahara, but he gives no explanation of this usage. According to M o n o d (in litt. 7.X. 1974), in parts of Mauritania, in Hodd, Azaouad, Timbuktoo, etc. 'Sahel' just means north. H e suggests that Chevalier, w h o reached the future Sahel zone from the south, adopted, perhaps inadvertently, the name of a
geographical direction for a botanical zone. It was a mere coincidence that the zone is situated along the southern fringes or 'coast* of the desert. A s applied in the present work it extends as a narrow east-west band, from Senegal to the Red Sea, where rainfall is 150-500 m m p.a. See Figures 1 andl\. S A V A N E STEPPIQUE, 173.
S A V A N N A , 45, 50ff, 113. This term, in the present work, is used only in a general sense for certain tropical landscapes in which both trees and grasses are conspicuous. N o attempt is made to use it in a precise classificatory sense. S A V O K A , 235-6. Malagasy secondary rain forest. S E B K H A . See S A B K H A
S H A T T (also transliterated C H O T T ) , 222. A classical Arabic term forriverbank or water's edge. In some modern dialects it is used as a synonym for 'sabkha'. 'Chott' is the form most frequently used in botanical literature. S P A R T E , 229. Lygeum spartum. SPEKBOOMVELD, 141,201. STEPPE, 45, 50, 149, 173. In this work the term steppe is not used for African vegetation. S U D D , 265, 266. The swamp region of the Upper Nile. Sometimes applied to similar swamps elsewhere. T A M P O K E T S A G R A S S L A N D , 239.
T A N D O S , 189. Seasonally flooded grassland on the coastal plain of Mozambique. TANETY GRASSLAND, 239.
TAPIA, 237. A vernacular n a m e applied to Uapaca bojeri and a forest type dominated by it. T E K E , 173. The Kalahari Sand-covered plateaux in Congo north of Brazzaville. V E L D . A n Afrikaans word used by South African botanists to signify vegetation. V L O E R E , 136, 139, 266. Brackish seasonal swamps in South Africa. W A D I . A desert valley, usually dry at the surface except after heavy rainfall. YAERE, 108. Grassland in the Chad basin subjected to prolonged flooding.
Geographical bibliography
The references cited below are not meant to be exhaustive, but include the more important works. In addition, m a n y of the publications mentioned in Chapters 3, 4, and 5 include information on vegetation. References to the offshore islands, other than Bioko, the Malagasy Republic, and Zanzibar, are given under the individual islands and archipelagos in Chapter X X I .
A L G E R I A (see also M A G H R E B and S A H A R A ) . Barry et al. (1970).
Barry & Faurel (1973). Cannon (1913). Guinet (1958). Guinochet & Quézel (1954). Hochreutiner (1904). Humbert (1928a). Killian (1961). Lemée (1953). Leredde (1957). Maire (1916). Monjauze (1958, 1968). Monjauze, Faurel & Schotter (1955). Ozenda (1954). Peyerimhoff (1941). Pons & Quézel (1955). Quézel (1954, 1956, 19576). Quézel & Santa (1962-63). R M & Schroter (1912). Simonneau (1954a, 1954e). A N G O L A . Airy Shaw (1947). Azancot de Menezes (1969). Barbosa (1970). Diruz (1973). Diniz & Aguiar (1969a, 19696, 1972, 1973). Gossweiler & Mendonça (1939, 1941). Humbert (1940a, 19406). Matos (1970). Matos & Sousa (1970). Mendes (1962). Mendonça (1961). Monteiro (1962, 1965, 1970). Redinha (1961). Teixeira (1968a, 19686). Teixeira & Correa de Pinho (1961). Teixeira & Matos (1967). Teixeira, Matos & Sousa (1967). Warburg (1903). Whellan(1965). B E N I N . Adjanohoun (1965, 1968). Aubréville (1937a). F A O (19806, 1980a> Paradis (1975a, 19756, 1976). Paradis, D e Souza & Houghnon (1978). B I O K O . A d a m s (1957). Guinea (1968). Mildbraed (19336). B O T S W A N A . Bawden (1965). Bawden & Stobbs (1963). Blair Rains & M c K a y (1968). Blair Rains & Yalala (1972). Bremekamp (1935). Campbell (1977). Cole & Brown (1976). Lang & Bremekamp (1935). Miller (1939, 1946). Pole Evans (1948a). Seagrief & D r u m m o n d (1958). Seiner (1911). Simpson (1975). Tinley (1966). Weare & Yalala (1971). Wild (19686). B U R U N D I . Deuse (1963, 1966). Lebrun (1956). Lewalle (1968, 1972, 1975). Pahaut & V a n der Ben (1962). Reekmans (1980a, 19806). Van der Ben (1961). C A M E R O U N . Aubréville (1948a). F A O (1980c, 1980a> Guillaume (1968). John B. Hall (1973). Hawkins & Brunt (1965). Hédin (1930). Jacques-Félix (1945, 1971). Keay (1955, 1959d). Ledermann (1912). Letouzey (1957, 1958, 1960, 1966, 1968a, 19686, 1969, 1975, 1977). Letouzey et coll. (1978, 1979). Maitland (1932). Mildbraed (1932, 1933a). Paulian & Gèze (1940). Portères (1946). Preuss (1892). Richards (1963a, 19636). Vaillant (1945). CENTRAL
AFRICAN
REPUBLIC.
Aubréville (1964). Chevalier
(1951). Guigonis(1968). Lanly (1966). Sillans(1951,1952a, 19526, 1952c, 1954, 1958).
C H A D (see also S A H A R A ) . Bruneau de Miré & Quézel (1959). Cavalho & Gillet (1960). Depierre & Gillet (1971). Gillet (1957, 1958, 1959a, 19596, 1960, 1961a, 19616, 1961c, 19626, 1963, 1964, 1968a, 19686). Grondard (1964). Maire & M o n o d (1950). Murat (1937). Pias (1970). Quézel (1958, 1959). Quézel, Bruneau de Miré & Gillet (1964). C O N G O . Begué (1965, 1967). Descoings (1972a). Farron (1968). Koechlin (1957, 1961). Makany (1972, 1976). Rollet (1963, 1964). D J I B O U T I . Chedeville (1972). Chevalier (1939). Verdcourt (1968). E G Y P T . Abdel R a h m a n & Batanouny (1959a, 19596, 1959c, 1966). Ayyad (1973, 1976). Ayyad & Elghonemey (1976). Batanouny (1964, 1966, 1973). Batanouny & Zaki (1973). Boulos (1966). Davis (1953). El Hadidi (1971). El Hadidi & Ayyad (1975). El Hadidi & Kosinova (1971). ElSharwaki & Fayad (1975). Hassib (1952). Kassas (1952, 1953a, 19536). Kassas & El-Abyad (1962). Kassas & Girgis (1964, 1965, 1970). Kassas & I m a m (1953, 1959). Kassas & Zahran (1962, 1965, 1967). Long (1955). Migahid (1947). Migahid et al. (1955-75). Stocker (1926, 1927). Tadros (1953, 1956). Tadros & Atta (1958a, 19586). E Q U A T O R I A L G U I N E A . Guinea (1946).
E T H I O P I A (including Eritrea). Beals (1968). Chaffey (1979). Cufodontis (1940). Engler (1906a). Gillett (1941). Hedberg (1971, 1975). H e m m i n g (1961). Klótzli (1975). Logan (1946). Mooney (1963). Negri (1913). Pichi-Sermolli (1938, 1939, 1940). Posnett & Reilly (1977). Scott (1952a, 19526, 1955,1958). Seebald (1972). G A B O N . Aubréville (1962, 1967a, 19676). Caballé (1978). Catinot (1978). Descoings (1974, 19766). Gloriod (1974). Hallé & Le Thomas (1968). Hallé, Le Thomas & Gazel (1967). Hladik (1974). Le Testu (1938). Saint-Aubin (1961, 1963). Villiers (1973). G A M B I A . Giglioli & Thornton (1965). Rosevear (1937). G H A N A . A h n (1958, 1959, 1961). Asare (1962). Aubréville (1959). Boughey (1957a). Brookman-Amissah et al. (1980). Chipp (1927). Douglas (1948). Foggie (1947a, 19476). Hall & Jenik (1968). Hall & Lawson (1972). Hall & Popple (1968). Hall & Swaine (1974, 1976, 1981). Jenik & Hall (1966, 1976). Jenik & Lawson (1968). Lawson (1956, 1968). Lawson, Armstrong-Mensah & Hall (1970). Lawson & Jenik (1967). Lawson, Jenik & Armstrong-Mensah (1968). Morton (1957). Ramsay & Rose Innes (1963). Swaine & Hall (1974). Taylor (1952, 1960). Vigne (1936). G U I N E A . A d a m (1947, 1948, 1950, 19586, 19686). A d a m & Jaeger (1976). Jaeger & A d a m (1947, 1950). Killian (1951). Killian & Schnell (1947). Lamotte, Aguesse & R o y (1962). Schnell (1950a, 1950c, 1950e, 1952a, 1952c, 1957, 1960,1961, 1968). G U I N E A - B I S S A U . Malato Beliz (1963). Malato Beliz & Alves Pereira (1965). Sousa (1958).
272
Geographical bibliography
I V O R Y C O A S T . Adjanohoun (1962, 1964, 1965). Adjanohoun & Aké Assi (1967). Adjanohoun, Aké Assi & Guillaumet (1968). Alexandre (1977, 1978). Aubreville (1957-58). Begué (1937). Bellier (1969). Bernard-Reversat, Huttel & Lemée (1978). César & Menault (1974). Chevalier (1908). Descoings (19726). Dugerdil (1970). Emberger, Mangenot & Miège (19506). Guillaumet (1967). Guillaumet & Adjanohoun (1971). Huttel (19756). Lamotte (1967, 1979). Lanly (1969). Latham & Dugerdil (1970). Mangenot (1950, 19556, 1971). Mangenot, Miège & Aubert (1948). Menaut & César (1979). Miège (1954, 1955, 1966). Monnier (1968). Paullian (1947). Poissonet & César (1972). Portères (1950). Roland (1967). Roland & Heydacker (1963). Rougerie (1957). Sarlin (1969). Schmidt (1973). Schnell (1957). Spichiger & Pamard (1973). Vuattoux(1968, 1970). K E N Y A . Agnew (1968). Ament (1975). Barkham & Rainy (1976). Birch (1963). Bogdan (1958). Coe (1967). Darling (19606). Edwards (1935, 1940). Faden (1970). Fries (1925). Fries & Fries (1948). Glover (1966). Glover & Trump (1970). Glover, Trump & Wateridge (1964). Glover & Wateridge (1968). Graham (1929). Greenway (1969). Hedberg (19696). Hemming (1972). Herlocker (1979a). Holland & Hove (1975). Isaac & Isaac (1968). Kerfoot (1964a). Lucas (1968). Mabberley (1975a). M o o m a w (I960). Sauer (1965). Staples & Hudson (1938). Synnott (1979a). Trapnell et al. (1966, 1969). Tweedie (1976). Verdcourt (n.d.). Wimbush (1937). L E S O T H O (see also S O U T H A F R I C A : A F R O M O N T A N E
REGION).
Bawden & Carroll (1968). LIBERIA. A d a m (1970, 1971a, 19716). Berger-Landefeldt (1959). Cooper & Record (1931). Jaeger & A d a m (1975). Kilnkel (1962, 1964, 1966a, 19666). Voerhoeve(1964,1968). L I B Y A (see also S A H A R A ) . Bartolo et al. (1977). Boulos (1972, 1975). Léonard (1969a, 1971). Nègre (1974). Scholz (1971). Thomas (1921). M A G H R E B (see also A L G E R I A , L I B Y A , M O R O C C O , and T U N I S I A ) .
Boudy (1948, 1950). Blaun-Blanquet (1928). Emberger (19556). Métro (1970). Quézel (1957a, 1976-77). Rikli (1943-48). M A L A G A S Y R E P U B L I C . Battistini & Richard-Vindard (1973). Dejardin, Guillaumet & Mangenot (1973). Granier (1979). Guillaumet & Koechlin (1971). Humbert (1927a, 19276, 19286, 1955a, 19556, 1959). Humbert & Cours Darne (1964-65). Keraudren (1968). Koechlin (1968, 1972). Koechlin, Guillaumet & Morat (1974). Leroy (1978). Morat (1973). Paulian et al. (1971, 1973). Perrier de la Bâthie (1921a, 1936). Rauh (1973). Segalen & Moureaux (1949). Straka (1960). Thomasson (1974,1976, 1977). M A L A W I . Brass (1953). Brown & Young (1974). Chapman (1962, 1968). Chapman & White (1970). Hall-Martin (1975). Hall-Martin & Fuller (1975). Howard-Williams (1975a, 1977). Howard-Williams & Walker (1974). Jackson (1954, 1968, 1969). Kalk, McLachlan & Howard-Williams (1979). M A L I R E P U B L I C . A d a m (1959). Audry & Rossetti (1962). Begué (1958). Duong-Huu-Thoi (1950a, 19506). Hagerup (1930). Jaeger (1950, 1956, 1959, 19656, 1968). Jaeger & Jarovoy (1952). Jaeger & Winkoun (1962). Raynal & Raynal (1961). Rossetti (1962). M A U R I T A N I A (see also S A H A R A ) . A d a m (1962c, 1965a, 1968c). Audry & Rossetti (1962). M o n o d (1952a, 1954a, 19546). Naegélé (1958a, 19586, 1959a, 19596, 1960). Roberty (1958). Sauvage (1946). M O R O C C O (see also M A G H R E B and
S A H A R A ) . Braun-Blanquet
(1928). Braun-Blanquet & Maire (1924). Cavassilas (1963). Dahlgren & Lassen (1972). Destremau (1974). Emberger (1925, 1932, 1936, 1939, 1948). Frodin (1923). Guinet & Sauvage (1954). Humbert (1924). Ionesco & Sauvage (1962, 1965-69). Ionesco & Stefanesco (1967). Killian (1941). Lecompte (1973). Maire (1924). Mathez (1973). Métro (1958). Nègre (1952a, 19526, 1953, 1956a, 19566, 1959). Nègre & Peltier (1976). Peltier (1971). Peyre (1973). Quézel (1952). Sauvage (1948, 1961, 1963, 1971). Theron & Vindt (1960). Vindt (1959). M O Z A M B I Q U E . Amico (1967). Barbosa (1952, 19686). G o m e s e Sousa (1967). Macedo (1970). Macnae & Kalk (1962a, 19626, 1969). Mendonça (1952). Myre (1960, 1962, 1964, 1971). Noel (1959). Pedro & Barbosa (1955). W U d (1953). N A M I B I A . Curson (1947). Dinter (1912, 1921). Giess (1962, 1968a, 19686, 1969, 1970, 1971). Giess & Tinley (1968). Keet (1950). Marloth (1909). Nordenstam (1970, 1974). Pearson (1907). Rennie (1936). Rutherford (1972). Tinley (1969, 1971). Volk (1966a, 19666). Volk & Leippert (1971). Walter (1936). Walter & Volk (1954). N I G E R (see also S A H A R A ) . Aubreville (19376, 1973). Collier & Dundas (1937). Dundas (1938). Fairbairn (1943). Peyre de Fabregues & Lebrun (1976). Pitot (1950a). L. P. White (1970). N I G E R I A . Adejuwon (1970, 1971a, 19716, 1971c). Ainslie (1926). Aubreville (1973). Bawden & Tuley (1966). Buxton (1935). Charter (1968). Charter & Keay (1960). Clayton (1957, 1958a, 19586, 1958c, 1961, 1963, 1966). Collier & Dundas (1937). Cook (1968). Fairbairn (1939). Forest Department, Nigeria (1948). Golding & G w y n n (1939). Hall, John B . (1971, 1977). Hall, John B. & Medler (1975a, 19756). Hall, John B . & Okali (1979). Hambler (1964). Hepper (1965, 1966). Hopkins (1962, 1965a, 19656, 1965a", 1966, 1968). Jackson (1964). Jones, A . D . P . (1950). Jones, E. W . (1950, 1955-56, 1963a, 19636). Keay (1947, 1948, 1949, 1951, 1952, 1959a, 1960, 1962, 1979). Keay & Onochie (1947). K e m p (1963). Kershaw (1968). Killick (1959). Kinako (1977). Lawton (1978a). MacGregor (1934). M o n o d (19526). Onochie (1961). Ramsay (1964). Ramsay & D e Leeuw (1964, 1965a, 19656). Redhead (1966). Richards (1939, 1957). Rosevear (1947, 1953, 1954). Ross (1954). Sanford (1968, 1969, 1974). Tuley (1966). Tuley & Jackson (1971). R W A N D A . Bouxin (1974, 1975a, 19756, 1975c, 1976). Deuse (1963, 1966, 1968). Frankart & Liben (1956). Hendrickx (1944). Lebrun (1955, 1956, 1961). Liben (1965). Renier (1954). Spinage (1972). Spinage & Guiness (1971, 1972). Troupin(1966). S A H A R A (see also individual countries). Bruneau de Miré & Quézel (1961). Capot-Rey (1953). Chipp (19306). Cloudsley-Thompson (1974). G r a m (1935). Guinea (1945, 1949). Kruger (1967). Lavauden (1927). Lebrun (1977, 1979). Léonard (1980). Maire (1933, 1938, 1940). Maire et al. (1925). Massart (1898). M o n o d (1938, 1958). Murât (1944). Ozenda (1958, 1977). Quézel (1965a, 1971). Quézel & Simonneau (1960, 1962). Schiffers (1971). Schulz (1979). Stocker (1926). Zolotarevsky & Murât (1938). S E N E G A L . A d a m (1953, 1956, 1958a, 1961a, 19616, 1961c, 1962a, 19626, 1964, 19656, 1968a, 1968¿). Aubreville (19486). Bule & Poupon (1972). Bourlière (1978). Devois (1948). Doumbia (1966). Jaeger (1949). Miège, Bodard & Carrère (1966). Miège, Hainard & Tchérémissinoff (1976). Naegélé (1959c). Pitot (19506). Pitot & A d a m (1954, 1955). Raynal, A . (1963). Raynal, J. (1964, 1968). Trochain(1940).
Geographical bibliography
SIERRA L E O N E . Cole (1967, 1968a, 19686, 1973). Cole & Jarrett (1969). Fox (1968a, 19686, 1968c, 1970). Gledhill (1963, 1970). Ifan-Dakar (1971). Jaeger (1965a, 1966, 1969, 1976). Jaeger & A d a m (1967, 1971, 1972). Jaeger, Lamotte & Roy (1966,1971). Jordan (1964). Morton (1968). S O M A L I R E P U B L I C . Bally (1968, 1976). Boaler & Hodge (1962, 1964). Ciferri (1939). Collenette (1931). Engler (1904). Gillett (1941). Gilliland (1952). Hemming (1965, 1966, 1968). Macfadyen (1950). Senni (1935). S O U T H A F R I C A (REPUBLIC OF)
G E N E R A L . Aco*cks (1953, 1964, 1971, 1975, 1977, 1979). Adamson (1938a, 19386). Aitken & Gale (1921). Bayer, Bigalke & Crass (1968). Bews (1912, 1913, 1916a, 19166, 1917a, 19176, 1918, 1925). Chipindall (1955). Coetzee & Werger (1975). Comins (1962). Dyer (1937). Edwards (1967). Goldblatt (1978). Hutchinson (1946). Killick (1968). Kruger (1979). Laughton (1937). Macnae (1963). Marloth (1887, 1908). Martin (1960a, 19606). Meredith (1955). Moll (1968c). Muir (1929). Phillips (1971). Pole Evans (1936). Roberts (1968). Rycroft (1968). Scheepers (1978). Scott (1951). Story (1952). Von Breitenbach (1972). Weintroub (1933). Wellington (1955). Werger (1978a, 1978c). West (1945, 1951). AFROMONTANE
REGION
(including the transition to the
Highveld). Granger & Schulze (1977). Herbst & Roberts (1974). Jacot Guillarmod (1962, 1963, 1968, 1969, 1971). Killick (1963, 1978a, 19786, 1978c, 1979). Moll (1966, 1968a, 1972a). Moll & Haigh (1966). Phillips (1928a, 19286, 1931a). Roberts (1961, 1966, 1969). Rycroft (1944). Taylor, H . C . (1962). Van Zinderen Bakker, E. M . , Jr (1971, 1973). Van Zinderen Bakker, E. M . , Sr (1955, 1965). Van Zinderen Bakker, E. M . , Sr & Werger (1974). C A P E R E G I O N . Adamson (1927, 1934, 1935, 1959). Boucher (1977, 1978). Boucher & Jarman (1977). Campbell & Moll (1977). Campbell, G u b b & Moll (1980). Day et al. (1979). Duthie (1929). Kruger (1977a, 19776, 1977c). Kruger & Taylor (1979). Marloth (1902, 1923, 1929). McLachlan, Moll & Hall (1980). Milewski (1977). Milewski & Esterhuysen (1977). Taylor (1953, 1961a, 1963, 1972a, 19726, 1977, 1978, 1979, 1980). Werger, Kruger & Taylor (1972a, 19726). KALAHARI-HIGHVELD
TRANSITION
ZONE.
Bredenkamp
(1975). Bredenkamp & Lambrechts (1979). Bredenkamp & Theron (1976, 1978, 1980). Leistner (1959, 1961a, 1967). Leistner & Werger (1973). Louw (1951). Mostert (1958). Potts & Tidmarsh (1937). Werger (1973a, 19736, 1978a). Werger & Coetzee (1977). Werger & Leistner (1975). K A R O O - N A M I B R E G I O N . Aco*cks (1964). Compton (1929a, 19296). Levyns (1950). Marloth (1909). Werger (19786). Werger & Coetzee (1977). TONGALAND-PONDOLAND
REGIONAL
MOSAIC.
Archibald
(1955). Bayer (1938). Bews (1920). Breen (1971). Downing (1980). Furness & Breen (1980). Henkel, Ballenden & Bayer (1936). Huntley (1965). Killick (1959). Martin (1965, 1966). Moll (19686, 1968a1, 19726, 1972c). Moll & Morris (1968). Moll & White (1978). Musil, Grunow & Bornman (1973). Penzhorn, Robertse & Olivier (1974). Rogers & Moll (1975). Varmeijer (1966). Van der Walt (1968). Venter (1976). Weisser (1978). Weisser & Marques (1979). Z A M B E Z I A N R E G I O N . Brynard (1964). Coetzee (1974, 1975). Coetzee et al. (1976). Galpin (1927). Gilliland (1962). Glover & Van Rensburg (1938). Grunow (1967). Schweickerdt (1933). Van der Meulen (1978, 1979). Van der Meulen & Westfall (1979). Verdoorn (1929). Wells (1964).
273
S U D A N . A d a m s (1967). Andrews (1945, 1948). Bari (1968). Begué (1958). Bruneau de Miré (1960). Bunting & Lea (1962). Chipp (1929, 1930a). Eyre, Ramsay & Jewitt (1953). G a y (1960). G a y & Berry (1959). Good (1924). Halwagy (1961, 1962a, 19626, 1963). Hanco*ck (1944). Harrison & Jackson (1958). Hunting Technical Services (1958,1964,1968). Jackson (1950,1951,1956). Jenkin et al. (1977). Jonglei Investigation T e a m (1954). Kassas (1956a, 19566, 1957). Lamprey (1975). M a h m o u d & Obeid (1971). Migahid (1947). Morison, Hoyle & Hope-Simpson (1948). Obeid & M a h m o u d (1971). Obeid & Seif el Din (1971a, 19716). Quézel (1969, 1970). Radwanski & Wickens (1967). Ramsay (1958). Ruxton & Berry (1960). Schweinfurth (1968). Smith (1949). Wickens (1977a). Wickens & Collier (1971). Willimot (1957). Worrall (1959, 1960, 19606). T A N Z A N I A (mainland). Albrecht (1964). Anderson & Herlocker (1973). Anderson & Talbot (1965). Backlund (1956). Bjornstad (1976). Boaler (1966). Boaler & Sciwale (1966). Brunnthaler (1914). Buchwald (1896). Burtt (1942). Clutton-Brock & Gillett (1979). Dean (1967). Engler (1894, 1900, 1903). Gillman (1949). Goetze & Engler (1902). Greenway (1933, 1955, 1965). Greenway & VeseyFitzGerald (1969). Herlocker (1975). Herlocker & Dirschl (1972). Jeffers & Boaler (1966). Kerfoot (19646). Klôtzli (1958). Lamprey (1963, 1964,1979). Leippert (1968). Milne (1947). Moreau (1935a). Pearsall (1957). Phillips (1930, 19316). Pielou (1952). Pitt-Schenkel (1938). Pócs (1974, 1976a, 19766, 1976c). Polhill (1968). Rodgers & H o m e w o o d (1979). Rodgers & Ludanga (1973). Salt (1951, 1954). Schmidt (1975a, 19756). Scott (1934). ToblerWolff & Tobler (1915). Vageler (1910). Vesey-FitzGerald (1955a, 1973a, 19746). Volkens (1897). Welch (1960). Welsh & Denny (1978). Werth (1915). W o o d (1965). T O G O . Aubréville (1937a). Busse (1907). E m (1979). F A O (1980a, 1980*0. TUNISIA. Burollet (1927). Gaussen & Vernet (1958). Knapp (19686). Lavauden (1928). Le Houérou (1959, 1962, 1967, 1969). Long (1954). Peyerimhoff (1941). Quézel & Bounaga (1975). Vanden Bergen (1977, 1979a, 19796,1980). U G A N D A . Bishop (1959). Buechner & Dawkins (1961). Dale (1954). Dawkins (1954). Denny (1971, 1973). Eggeling (1935, 1938, 1947). Harrington & Ross (1974). Jackson & Gartlan (1965). Kerfoot (1965). Lang Brown & Harrop (1962). Langdale-Brown (1959a, 19596, 1960a, 19606, 1960c). Langdale-Brown, Osmaston & Wilson (1964). Laws (19706). Leggat (1965). Lind (1956a, 19566). Lind & Visser (1962). Lock (1973, 1977a, 19776). Loveridge (1968). Osmaston (1968). Ross (1955a, 19556). Snowden (1933, 1953). Thomas, A . S. (1941, 1943, 1945, 1946). Wilson (1962). W o o d (1960). Z A I R E . Aubréville (1957). Balle (1953). Bamps (1975). Bernard (1945). Bouillenne, Moureau & Deuse (1955). Bourbeau et al. (1955). Bourguignon, Streel & Calembert (1960). C h a m b ó n & Leruth (1954). Colonval-Elenkov & Malaisse (1975). Compère (1970). Cornet D'Elzius (1964). Delevoy (1933). Delevoy & Robert (1935). Delvaux (1958). Demaret (1958). Denisoff & Devred (1954). D e Saeger (1954). Desenfans (1950). Deuse (1960). Devred (1956, 1957, 1958). Devred, Sys & Berce (1958). D e Wildeman (1932, 1934). Dieterlen (1978). Diels (1915). Dubois (1955). Duvigneaud (1949a, 19496, 1950, 1952, 1953, 1958, 1959). Duvigneaud & Denaeyer-de Smet (1960, 1963). Duvigneaud & Symoens (1951). Evrard (1957, 1965, 1968). Focan & Mullenders (1949, 1955). Frankart & Liben (1956). Fresón, Goffinet & Malaisse (1974). Gérard (1960).
274
Geographical bibliography
Germain (1945, 1949, 1952, 1965, 1968). Germain, Croegaert & Sys (1955). Germain & Evrard (1956). Gilson et al. (1957). Gilson, Van Wambeke & Gutzwiller (1956). Hauman (1933). Hendrickx (1944, 1946). Holowaychuk et al. (1954). Jongen et al. (1960). Lebrun (1935, 1936a, 19366, 1942, 1947, 1954, 1955, 1957, 1959, 1960a, 19606, 1960a", 1968, 1969). Lebrun & Gilbert (1954). Léonard, A . (1959, 1962). Léonard, J. (1947, 1950, 1951, 1952a, 19526, 1953, 1954). Liben (1958, 1962). Louis (1947a, 19476, 1947c). Malaisse (1975, 1976a, 19766). Malaisse & Anastassiou-Socquet (1977). Malaisse & Grégoire (1978). Meessen (1951). Mullenders (1953, 1954, 1955). Nanson & Gennart (1960). Pahaut & Van der Ben (1962). Pecrot & A . Léonard (1960). Peeters (1964). Pierlot (1966). Pynaert (1933). Robyns (1932, 1936, 1937, 1941, 1948a, 19486, 1950). Schmitz (1950, 1952a, 19526, 1962, 1963a, 19636, 1971, 1977). Streel (1962, 1963). Symoens (1953, 1963). Symoens & Ohoto (1973). Sys & Schmitz (1959). Taton (1949a, 19496). Taton & Risopoulos (1955). Thomas (1941). Van der Ben (1959). Vanderyst (1932, 1933). Van Meel (1952, 1953, 1966). Van Wambeke & Evrard (1954). Van Wambeke & Liben (1957). Z A M B I A . Astle (1965a, 19656, 1969). Astle, Webster & Lawrance (1969). Balon & Coche (1974). Boughey (1964). Cole (1963a). Cottrell & Loveridge (1966). Debenham (1952).
Drew & ReUly (1972). Edmonds (1976). Fanshawe (1961, 1968, 1969). Fanshawe & Savory (1964). Fries (1913, 1915, 1921). Horscroft (1961). Kornas (1977, 1978, 1979). Lawton (1963, 1964, 1967a, 19676, 1972, 19786). Martin (1940, 1941). Mitchell (1969). Seagrief (1962). Trapnell (1953, 1959). Trapnell & Clothier (1937). Trapnell et al. (1976). Trapnell, Martín & Allan (1950). Verboom (1965, 1966). Verboom & Brunt (1970). Vesey-FitzGerald (1955a). White (1968). Z A N Z I B A R . Robins (1976). Werth (1901). Z I M B A B W E . Anderson & Walker (1974). Atwell (1970). Banks (1976). Barclay-Smith (1964). Boughey (1961, 1963a, 19636). Crook (1956). Dye & Walker (1980). Farrell (1968a, 19686). Gilliland (1938). Goldsmith (1976). Goodier & Phipps (1961, 1962). G u y (1977). Henkel (1931). Ingram (1960). Jacobsen (1967, 1968, 1970, 1973). Kelly & Walker (1976). Kennan (1972). Kennan, Staples & West (1955). Lang (1952). Magadza (1970). Mitchell, B. L. (19616). Mitchell, D . S. (1969). Phipps & Goodier (1962). Proctor & Craig (1978). Rattray (1957, 1961). Rattray & Wild (1955). Strang (1974). Thomas, Walker & Wild (1977). Werger, Wild, & Drummond (1978a, 19786). West (1958). Wild (1952a, 1953, 1955, 19646, 1965, 1968c, 1968a1, 1968e, 1970, 1974a, 19746, 1974c, 1974a", 1974e, 1975).
Alphabetical bibliography
Abbreviations employed: AETFAT
Association pour l'Étude Taxonomique de la Flore d'Afrique Tropicale/ Association for the Taxonomic Study of the African Flora ASGA Association des Services Géologiques Africains/ Association of African Geological Surveys CCTA Commission de Coopération Technique en Afrique au Sud du Sahara/Commission for Technical Co-operation in Africa South of the Sahara CNRS Centre National de la Recherche Scientifique (15 Quai Anatole-France, 75700 Paris, France) CSA Conseil Scientifique pour l'Afrique au Sud du Sahara/ Scientific Council for Africa South of the Sahara FAO Food and Agriculture Organization of the United Nations/ Organisation des Nations Unies pour l'Alimentation et l'Agriculture (Rome, Italy) F U L R E A C Fondation de l'Université de Liège pour les Recherches en Afrique Centrale IEMVT Institut d'Élevage et de Médecine Vétérinaire des Pays Tropicaux (10 R u e Pierre-Curie, 94700 Maisons-Alfort, France) IFAN Institut Français d'Afrique Noire (up to 1966), Institut Fondamental d'Afrique Noire (after 1966) (Dakar, Senegal) INEAC Institut National pour l'Étude Agronomique du Congo (Publications obtainable from S E R D A T , q.v.) IPAL Integrated Project on Arid Lands/Projet Intégré sur les Terres Arides IUCN International Union for Conservation of Nature and Natural Resources (Morges, Switzerland) MAB M a n and the Biosphere Programme (of Unesco) ORSTOM Office de la Recherche Scientifique et Technique Outre-Mer (70-74 Route d'Aulnay, 93 Bondy, France) SERDAT Service de Documentation en Agronomie Tropicale (Rue Defacqz 1, 1050 Brussels, Belgium) UNDP United Nations Development Programme UNESCO United Nations Educational, Scientific and Cultural Organization/ Organisation des Nations Unies pour l'Éducation, la Science et la Culture (7 Place de Fontenoy, 75700 Paris, France)
ABDEL R A H M A N , A. A.; BATANOUNY, K. H. 1959a. Seasonal variations in the desert vegetation along Cairo-Suez road. Bull. Inst. Désert Egypte, 9, p. 1-10. ; . 1959¿. The phenology of the desert vegetation in relation to environment. Bull. Inst. Désert Egypte, 9, p. 1119. ; . 1959c. Root development and establishment of plants under desert conditions. Bull. Inst. Désert Egypte, 9, p. 41-50. ; . 1965. Vegetation and root distribution in the different microhabitats in wadi Hof. Bull. Inst. Désert Egypte, 15, p. 55-66. A C O C K S , J. P. H . 1953. Veld types of South Africa. Mem. bot. Surv. S. Afr., 28, p. 1-192, with coloured vegetation m a p 1:1500000. . 1964. Karoo vegetation in relation to the development of deserts. In: Davis, D . H . S. (éd.), p. 100-12. . 1971. T h e distribution of certain ecologically important grasses in South Africa. Mitt. Bot. Staatssamml. Munch., 10, p. 149-60. . 1975. Veld types of South Africa. 2nd ed. Mem. bot. Surv. S. Afr., 40, p. 1-128, with coloured vegetation m a p 1:1500000. . 1977. Riverine vegetation of the semi-arid and arid regions of South Africa. J. S. Afr. biol. Soc, 17, p. 21-35. . 1979. The flora that matched the fauna. Bothalia, 12, p. 673-709. A D A M , J. G . 1947. L a végétation de la région de la source du Niger. Annls Géogr., 56, p. 192-200. . 1948. Les reliques boisées et les essences des savanes dans la zone préforestière en Guinée française. Bull. Soc. bot. Fr., 95, p. 22-6. . 1950. Les formations végétales ligneuses secondaires de Guinée française. Conf. int. Afr. OccicL em Bissau, 1947, 2 (la), p. 2 2 5 ^ 1 . . 1953. Notes sur la végétation des Niayes de la presqu'île du C a p Vert (Dakar, A O F ) . Bull. Soc. bot. Fr., 100, p. 153-8. . 1956. L a végétation de l'extrémité occidentale de l'Afrique. L a pointe des Almadies aux environs de Dakar (Sénégal). Bull. IFAN, sér. A , 18, p. 685-702. . 1958a. Flore et végétation de la réserve botanique de Noflaye (environs de Dakar, Sénégal). Bull. IFAN, sér. A , 20, p. 809-68. . 19586. Éléments pour l'étude de la végétation des hauts plateaux du Fouta Djalon (Secteur des Timbis), Guinée française. I. La flore et ses groupements. Dakar, G o u vernement Général de l ' A O F , Bureau des Sols. 80 p., with coloured vegetation m a p 1:50000. . 1959. Contribution à l'étude floristique des pâturages du Soudan français. In: Charreau, C , et al. (eds.), Études des
276
Alphabetical bibliography
pâturages tropicaux de la zone soudanienne, p. 49-75, with map. . 1961a. L a végétation d u bois sacré d'Oussouye (Casamance) et quelques intrusions du domaine de la forêt dense en basse Casamance. Bull. IFAN, sér. A , 23, p. 1-10. . 19616. Florule et végétation de la grande Mamelle de Dakar (Phare). Bull. IFAN, sér. A , 23, p. 406-22. . 1961c. Flore et végétation de l'île de la Madeleine (Dakar). Bull. IFAN, sér. A , 23, p. 708-15. . 1962a. Contribution à l'étude de la flore et de la végétation de l'Afrique occidentale. La Basse-Casamance (Sénégal). Bull. IFAN, sér. A , 24, p. 116-53. . 19626. Éléments pour l'étude des groupements végétaux de la presqu'île du Cap-Vert (Dakar). La série du massif de N'Diass. Bull. IFAN, sér. A , 24, p. 154-67. . 1962c. Itinéraires botaniques en Afrique occidentale; flore et végétation d'hiver de la Mauritanie occidentale; les pâturages; inventaire des plantes signalées en Mauritanie. J. Agrie, trop. Bot. appl, 9, p. 85-200. . 1964. Contribution à l'étude de la végétation du lac de Guiers (Sénégal). Bull. IFAN, sér. A , 26, p. 1-72. . 1965a. L a végétation du delta du Sénégal en Mauritanie. Bull. IFAN, sér. A , 27, p. 121-38, with small vegetation map. . 19656. GénéraUtés sur la flore et la végétation du Sénégal. Étud. Sénégal, 9 (3), p. 155-214. . 1968a. L a flore et la végétation du Parc National du Niokolo-Koba (Sénégal). Adansonia, sér. 2, 8, p. 439-59. . 19686. Flore et végétation de la lisière de la forêt dense en Guinée. Bull. IFAN, sér. A , 30, p. 920-52, with vegetation m a p 1:25000. . 1968c. L a Mauritanie. In: Hedberg, I.; Hedberg, O . (eds.), p. 49-51, with small vegetation m a p . . 1968a". Sénégal. In: Hedberg, I.; Hedberg, O . (eds.), p. 65-9. . 1970. État actuel de la végétation des monts N i m b a au Libéria et en Guinée. Adansonia, sér. 2,10, p. 193-211. . 1971a. L a végétation littorale aux environs de Buchanan (Libéria). Bull. IFAN, sér. A , 32, p. 995-1018. . 19716. Aperçu sur la flore et la végétation des Monts N i m b a au Libéria. In: Flore descriptive des Monts Nimba, 1. Mém. Mus. natn. Hist, nat., Paris, n.s., sér. B (Bot.), 20, p. 23-144. ADAM,
J. G . ; J A E G E R , P . 1976.
Suppression de la floraison
consécutive à la suppression des feux dans les savanes et prairies de la Guinée (Afrique occidentale). C.r. hebd Séanc. Acad. Sci., Paris, 282, p. 637-9. A D A M S , C . D . 1957. Observations on the fernfloraof Fernando Po. I. A description of the vegetation with particular reference to the Pteridophyta. J. Ecol, 45, p. 479-94. A D A M S , M . E . 1967. A study of the ecology of Acacia mellifera, A. seyal, and Balanites aegyptiaca in relation to land clearing. / . appl. Ecol., 4, p. 221-37. A D A M S O N , R . S. 1927. The plant communities of Table Mountain: preliminary account. / . Ecol., 15, p. 278-309. . 1934. The vegetation andfloraof Robben Island. Trans. R. Soc. S. Afr., 22, p. 279-96. . 1935. The plant communities of Table Mountain. III. A six years' study of regeneration after burning. / . Ecol., 23, p. 44-55. . 1938a. The vegetation of South Africa. London, British E m p . Veg. C o m m . 235 p., with 3 small vegetation maps. . 19386. Notes on the vegetation of the Kamiesberg. Mem. bot. Surv. S. Afr., 18, p. 1-25.
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Index of plant names
Important synonyms are included in this index, but no attempt is m a d e to indicate which are 'correct'. This is often a matter of opinion and sometimes of controversy. Page references are given after the n a m e which is 'preferred' by the author. It should not be inferred, however, that the preferred n a m e will prove to be taxonomically 'correct' or otherwise generally acceptable. S o m e names are preferred because they are so well k n o w n in the ecological literature that a wise society would choose to conserve them. S o m e names, which are not preferred, might eventually prove to be taxonomically acceptable, but they have been proposed in incomplete or otherwise controversial revisions, and at present there is no guarantee that they are here to stay. Synonyms in the index are cross-referenced to the preferred n a m e . In the text, synonymy is usually given only where the n a m e is first mentioned, or in connection with important source materials where the names are used.
Abies alba Miller (Pinaceae), 156 A. numidica Delannoy ex Carrière, 60, 146, 149, 150, 152, 155,156 A. pinsapo Boiss., 60, 146, 155, 156 ss. marocana (Trabut) Ceballos, 149, 150, 152,156 Abrus precatorius L. (Leguminosae: Papilionoideae), 206 Abutilón (Malvaceae), 220 A. fruticosum Guill. & Perr., 222 Acacia (Leguminosae: Mimosoideae), 31, 62, 63, 64, 65, 89, 98, 103, 107, 108, 110, 111, 113, 115, 116, 125, 128, 144, 181,182, 191, 201, 206, 216, 219, 220, 241, 242, 254 A. albida Del., 28, 55, 91, 95, 105, 141, 144, 203, 208, 209, 210, 211, 212, 214, 215, 220, 222, 251, 252 A. ataxacantha D C , 90, 209, 211 A. borleae Burtt Davy, 201 A. brevispica H a r m s , 129 A. burkei Benth., 201 A. bussei H a r m s ex Sjóstedt, 114, 116,188 A. caffra (Thunb.) Willd., 96, 196, 201 A. clavigera E . Meyer: see A robusta ss. clavigera A. cyanophylla Lindley, 135, 231 A. cyclops A . C u n n . ex G . D o n , 135 A.davyiN.E.Bi.,96,196,201 A. drepanolobium H a r m s ex Sjóstedt, 52, 115, 116,121,128 A. dudgeonii Craib ex Holland, 105 A. ehrenbergiana Hayne, 207, 219, 222 A. elatior Brenan, 117 A. erioloba E . Meyer, 31, 90, 95, 97, 137, 140, 141, 144, 191, 193,194 A. erubescens W e l w . ex Oliver, 95 A. etbaica Schweinf., 116, 120, 121 A. farnesiana (L.) Willd., 252 A. flava (Forssk.) Schweinf. not of Spreng. ex D C : see A. ehrenbergiana
A.fleckiiSchinz,90, 193 A. galpinii Burtt Davy, 91 A. gerrardii Benth, 96, 128, 129, 182, 201, 209 A. gillettiae Burtt Davy: see A. luederitzii A. giraffae Willd.: see A. erioloba A. gourmaensis A . Chev., 105 A. gummifera Willd., 149, 225, 226, 227, 228, 229, 231 A. haematoxylon Willd., 191, 194 A. hebeclada D C , 191, 193 A. hereroensis Engl., 193 A. heterophylla Willd., 258 A. hockii D e Wild., 87, 105,115, 128, 129, 174, 182 A. hórrida (L.) Willd., 120 A. kamerunensis Gandoger, 81 A. karroo Hayne, 137, 141, 159, 193, 194,195,201 A.kirkii Oliver, 95,115 ss. mildbraedii (Harms) Brenan, 181,182 A. laeta R . Br. ex Benth., 206, 207, 212, 222 A. lahai Steud. & Hochst. ex Benth., 130 A. luederitzii Engl., 96, 193 A. macrostachya Reichenb. ex D C , 105 A. macrothyrsa H a r m s , 105 A. malacocephala H a r m s , 116 A. melanoxylon R . Br., 135 A. mellifera (Vahl) Benth., 108, 114, 116, 120, 121, 126, 128, 188, 191, 193, 203, 207, 208, 209, 210, 212, 213,214 ss. detinens (Burchell) Brenan, 137, 140 A. montis-usti M e r x m . & A . Schreiber, 140 A. nebrownii Burtt Davy, 267 A. nigrescens O U ver, 94, 95, 201 A. nilotica (L.) Willd. ex Del., 96,128, 188, 252 ss. adansonii (Guill. & Perr.) Brenan: see ss. adstringens ss. adstringens (Schumach. & Thonn.) Roberty, 105, 209, 210, 220 ss. kraussiana (Benth.) Brenan, 196, 267 ss. nilotica, 108 ss. subalata (Vatke) Brenan, 114, 121, 129 A. nubica Benth., 121, 208, 210, 212 A. pennivenia Balf.f., 255 A. permixta Burtt Davy, 96 A. polyacantha Willd., 63, 89, 128,129, 172, 209 ss. campylacantha (Hochst. ex A . Rich.) Brenan, 91, 95, 105, 174 A. pseudofistula H a r m s , 116 A. redacta J. H . Ross, 140 A. reficiens W a w r a , 120,121 ss. misera (Vatke) Brenan, 114, 120 ss. reficiens, 145, 193, 194 A. rehmanniana Schinz, 96 A. robusta Burchell, 196, 201
326
Index ofplant names
Acacia (Leguminosae: Mimosoideae)—contd ss. clavigera (E. Meyer) Brenan, 91, 95, 129 ss. robusta, 96 ss. usambarensis (Taubert) Brenan, 117, 128, 188 A. robynsiana M e r x m . & A . Schreiber, 140 A. Senegal (L.) Willd., 105, 107, 120, 128, 182, 189, 201, 203,206,207,208,210,211, 212,213 var. kerensis Schweinf., 121 A. seyal Del., 62, 105, 107, 108, 115, 116, 120, 121, 128, 203, 209, 210, 211, 212, 214, 222 A. sieberana D C , 95, 96, 105, 107, 128, 129, 174, 196, 201, 209,210,211 A. stenocarpa Hochst. ex A . Rich.: see A. seyal A. tanganyikensis'ñxenaxi, 116 A. tenuispina I. Verdoorn, 96 A. thomasii H a r m s , 114 A. tortilis (Forssk.) Hayne, 31, 91, 94, 95, 96, 113, 114, 116, 117, 120, 121, 127, 128, 145, 193, 206, 207, 208, 212,213,218,219,220,221,224 ss. raddiana (Savi) Brenan, 219 A. welwitschii Oliver, 90 A. xanthophloea Benth., 30, 31, 91,128,129, 267 A. zanzibarica (S. Moore) Taubert, 189 Acalypha chirindica S. Moore (Euphorbiaceae), 90, 98 Acanthosicyos horridus W e l w . ex Hook.f. (Cucurbitaceae), 144, 145 Acanthospermum hispidum D C . (Compositae), 252 Acanthus (Acanthaceae), 158 A. eminens C . B . Clarke, 122 A. mollis L., 149 Acer campestre L . (Aceraceae), 147, 149, 156 A. granatense Boiss., 156 A. monspessulanum L., 149,151, 156,157 A. obtusatum Waldst. & Kit. ex Willd., 156 Aceras (Orchidaceae), 158 Achyranthes áspera L . (Amaranthaceae), 90, 182 Aco*kanthera (Apocynaceae), 115 A. schimperi ( A . D C . ) Oliver, 115 Acridocarpus excelsus Adr. Juss. (Malpighiaceae), 243 A. smeathmannii ( D C . ) Guill. & Perr., 81 Acrocephalus sericeus Briq. (Labiatae), 100 Acroceras macrum Stapf (Gramineae), 100 Acrostichum aureum L. (Pteridaceae), 188, 262, 263 Actiniopteris (Actiniopteridaceae), 238 Adansonia (Bombacaceae), 241, 242 A. digitata L., 31, 62, 90, 94, 95, 96, 114, 116, 128, 174, 189, 212,213 A. fony Bâillon, 241, 242 A. grandidieri Bâillon, 241 A. madagascariensis Bâillon, 255 A. rubrostipa Jumelle & Perrier, 241 A. za Bâillon, 241,242 Adenia (Passifloraceae), 242 A. globosa Engl., 114, 188 A. pechuellii (Engl.) H a r m s , 142 A. venenata Forssk., 113 Adenium multiflorum Klotzsch (Apocynaceae): see A obesum A. obesum (Forssk.) R o e m . & Schult., 94,114 A. socotranum Vierh., 115, 155, 255 Adenocarpus bacquei Battand. & Pitard (Leguminosae: Papilionoideae), 153 A. foliolosus (Aiton) D C , 249 A. viscosus W e b b & Berth., 248 Adenolobus (Leguminosae: Caesalpinioideae), 137 A. garipensis (E. Meyer) Torre & Hillcoat, 140
A. pechuelii (Kuntze) Torre & Hillcoat, 140,144 Adiantum capillus-veneris L . (Adiantaceae), 224 A. vogelii Mett. ex Keys., 75 Adina microcephala (Del.) Hiern (Rubiaceae), 91,105 Aeluropus lagopoides (L.) Trin. ex Thwaites (Gramineae), 230 A. repens (Desf.) Pari.: see A. lagopoides Aeonium (Crassulaceae), 246, 249 A. arboreum (L.) W e b b , 226, 228 Aerva javanica (Burm.f.) Juss. ex Schultes (Amaranthaceae), 115 A. pérsica (Burm.f.) Merr., 221, 252 A. tomentosa Forssk.: see A. javanica Aeschynomene elaphroxylon (Guill. & Perr.) Taub. (Leguminosae: Papilionoideae), 266 A. pfundii Taub., 266 A. trigonocarpa Taub., 98 Aframomum angustifolium K . Schum. (Zingiberaceae), 236 A. biauriculatum K . Schum., 96 Afrobrunnichia (Polygonaceae), 74 Afrocrania (Cornaceae), 162 A. volkensii (Harms) Hutch., 167 Afrormosia angolensis (Baker) D e Wild.: see Pericopsis angolensis A. elata H a r m s : see P. elata A. laxiflora Benth. ex Baker: see P. laxiflora Afrosersalisia cerasifera (Welw.) Aubrév. (Sapotaceae), 187 Afrotrilepis pilosa (Boeckeler) J. Raynal (Cyperaceae), 81 Afzelia africana Smith (Leguminosae: Caesalpinioideae), 79, 80, 83, 85, 105, 107, 178 A. bipindensis H a r m s , 74 A. quanzensis W e l w . , 93, 95, 96, 97, 99, 117, 187, 188, 189, 200, 201 Agathophora (Chenopodiaceae), 218 Agathosma (Rutaceae), 132 Agauria (Ericaceae), 237 A. buxifolia (Lam.) Cordem., 258 A. salicifolia (Lam.) Hook.f. ex Oliver, 237, 238, 258 Agave (Agavaceae), 159 Agelaea (Connaraceae), 75, 253 Ageratum conyzoides L . (Compositae), 252 Agrostis (Gramineae), 169 A. azorica (Hochst.) Tutin & E . W a r b . , 246 A. elliotii Hackel, 239 Aichryson (Crassulaceae), 246 Aizoanthemum dinteri (Schinz) Friedrich (Aizoaceae), 142 Aizoon (Aizoaceae), 266 A. canariense L., 229 A. dinteri Schinz: see Aizoanthemum dinteri A. mossamedense W e l w . ex Oliver, 144 A. virgatum W e l w . ex Oliver, 144 Ajuga (Labiatae), 236 A. ophrydis Burchell ex Benth., 194 Alangium chínense (Lour.) H a r m s (Alangiaceae), 181, 187 Alberta (Rubiaceae), 237 A. minor Bâillon ex K . Schum., 237, 238 Albizia (Leguminosae: Mimosoideae), 98,181, 241 A. adianthifolia (Schumach.) W . F . Wight, 74, 83, 173, 178, 187, 199 A. amara (Roxb.) Boivin, 95, 106, 116, 208, 209, 210, 212, 213,214 A. anthelmintica Brongn., 188, 191,193,194, 212 A. antunesiana H a r m s , 96, 97 A. aylmeri Hutch., 210 A. brachycalyx Oliver: see A. petersiana A. chevalieri H a r m s , 105, 107 A. falcata (L.) Back, ex Merr., 257
Index of plant names
A. ferruginea (Guill. & Perr.) Benth., 81, 178 A. forbesii Benth., 200 A. glaberrima (Schumach. & Thonn.) Benth., 117 A. grandibracteata Taubert, 182 A. gummifera (J. F . Gmelin) C . A . Smith, 130, 166,181 A. harveyi Fourn., 95,116, 129 A. lebbeck (L.) Benth., 257 A. malacophylla (Steud. ex A . Rich.) Walp., 211 A. petersiana (Bolle) Oliver, 97,188 A. sericocephala Benth.: see A amara A. tanganyicensis Baker f., 96 A, versicolorWelw. ex Oliver, 91, 95,173,174, 201 A. zimmermannii Harms, 117 A. zygia ( D C . ) J. F. Macbr., 83,105, 172, 178, 211 Alchemilla (Rosaceae), 236 Alchornea cordifolia (Schumach. & Thonn.) Muell. Arg. (Euphorbiaceae), 83 A. occidentalis (Muell. Arg.) Pax & K . Hoffm., 90 Allanblackia stuhlmannii (Engl.) Engl. (Guttiferae), 187 Allium (Alliaceae), 158 Allmaniopsis (Amaranthaceae), 111 Allophylus abyssinicus (Hochst.) Radlk. (Sapindaceae), 122 A. africanus P. Beauv., 182 Alloteropsis cimicina (Retz.) Stapf (Gramineae), 213 A. semialata (R. Br.) Hitchco*ck, 194, 202, 239 Alluaudia (Didiereaceae), 242 A. ascendens Drake, 242 A. procera Drake, 242 Alluaudiopsis (Didiereaceae), 242 Alnus glutinosa (L.) Gaertner (Betulaceae), 147,149, 152,156 Aloe(Liliaceae), 111, 114, 115, 116, 129, 137, 140, 174, 198, 201, 238, 242 A. arborescens Miller, 96, 196 A. asperifolia Berger, 142 A. bainesii Dyer, 200 A. ballyi Reynolds, 117 A. breviscapa Reynolds & Bally, 116 A. candelabrum Berger, 201 A. capitata Baker var. cipolinicola H . Perrier, 243 A. dichotoma Masson, 140 A. eminens Reynolds & Bally, 115 A / e r o x Miller, 135,195,201 A. kedongensis Reynolds, 115 A. littoralis Baker, 145 A. marlothii Berger, 96, 201 A. perryi Baker, 115, 255 A. pillansii L . Guthrie, 140 A. plicatilis (L.) Miller, 134 A. rigens Reynolds & Bally, 116 A. scobinifolia Reynolds & Bally, 116 A. speciosa Baker, 137, 201 A. spectabilis Reynolds, 201 A. volkensii Engl., 129 Alstonia boonei D e Wild. (Apocynaceae), 81, 181 A. congensis Engl., 82 Alyssum serpyllifolium Desf. (Cruciferae), 230 A. spinosum L . , 158 Amanoa bracteosa Planchón (Euphorbiaceae), 82 Amaranthus graecizans L. (Amaranthaceae), 212 Amblygonocarpus andongensis (Welw. ex Oliver) Exell (Leguminosae: Mimosoideae), 87, 96, 97,105 Amelanchier ovalis Medicus (Rosaceae), 158 Ammannia gracilis Guill. & Perr. (Lythraceae), 204 Ammodaucus (Umbelliferae), 221
327
Ampelodesma mauretanicum (Poir.) Th. Durand & Schinz (Gramineae), 149,154, 155,157, 158, 159,229 y4mpAi'was(Leguminosae: Caesalpinioideae), 74 Anabasis aphylla L. (Chenopodiaceae), 218, 229, 266 A. aretioides M o q . & Coss.: see Fredolia aretioides A. articulata (Forssk.) M o q . , 221, 222, 224 A. oropediorum Maire, 230 Anacampseros (Portulacaceae), 137, 140 A. albissima Marloth, 142 Anacamptis (Orchidaceae), 158 Anacardium (Anacardiaceae), 189 A. occidentale L., 174 Anadelphia afzeliana (Rendle) Stapf (Gramineae), 84 A. leptocoma (Trin.) Pilger, 84 A. trispiculata Stapf, 84 Anagyris (Leguminosae: Papilionoideae), 147 A.foetidaL., 158 Anastatica (Cruciferae), 218 A. hierochuntica L., 221 A ñas trabe (Scrophulariaceae), 199 A. integerrima E . Meyer ex Benth., 199, 200 Ancistrophyllum (Palmaceae), 82 Androcymbium (Liliaceae), 221 Andropogon (Gramineae), 169 A. amplectens Nées, 196 A. appendiculatusNees, 194 A. brazzae Franchet, 100 A. canaliculatus Schumach., 178 A. curvifolius W . D . Clayton, 84 A. distachyos L., 208, 211 A. eucomus Nées, 239 A. gayanus Kunth, 85, 97,108, 206, 209, 210 A. greenwayi Napper, 126,127 A. kelleri Hacktl, 116 A. perligulatus Stapf, 84 A. schirensis Hochst. ex A . Rich., 85, 100, 101, 173, 194, 196 A. tectorum Schumach. & Thonn., 83, 84, 85 A. trichozygus Baker, 239 Androstachys (Euphorbiaceae), 87 A.johnsonii Prain, 96 Aneilema johnstonii K . Schum. (Commelinaceae), 90 Aneulophus (Linaceae), 74 Angkalanthus (Acanthaceae), 111 Angraecum (Orchidaceae), 238 Aningeria adolfi-friedericii (Engl.) Robyns & G . Gilbert (Sapotaceae), 85, 164, 187 A. altissima (A. Chev.) Aubrév. & Pellegr., 74, 79, 90, 181 A. pseudoracemosa J. H . Hemsley, 186 A. robusta (A. Chev.) Aubrév. & Pellegr., 79 Anisophyllea boehmii Engl. (Rhizophoraceae), 97 A. cabote Henriq., 253 A. gossweileri Engl. & v. Brehm., 173 A. pomífera Engl. & v. Brehm., 93 A. quangensis Engl., 173 Annona senegalensis Pers. (Annonaceae), 83, 85, 105, 107, 174, 189 Anogeissus (Combretaceae), 62,106 A. leiocarpus ( D C . ) Guill. & Perr., 55, 80, 83, 105, 106, 107, 203,208,209,210,211,214 Anonidium usambarense R . E . Fries (Annonaceae), 187 Anopyxis (Rhizophoraceae), 74 Ansellia gigantea Reichb.f. (Orchidaceae), 93, 201 A. nilotica (Baker) N . E . Br.: see A. gigantea Anthephora argéntea Goossens (Gramineae), 191, 193
328
Index ofplant names
Anthephora (Gramineae)—coníd A. lynesii Stapf & C . E . Hubbard, 209, 210 A. pubescens Nees, 193, 194 A. schinzü Hackel, 143 Anthocleista (Loganiaceae), 80 A. nobilis G . D o n , 83 A. schweinfurthii G ü g , 91, 181 Anthonotha (Leguminosae: Caesalpinioideae), 74 A. obanensis (Baker f.) J. Léonard, 81 A. pynaertii (De Wild.) J. Léonard, 181 Anthospermum rigidum Eckl. & Zeyh. (Rubiaceae), 194 Anthostema (Euphorbiaceae), 235 A. aubryanum Bâillon, 253 Anthoxanthum madagascariense Stapf (Gramineae), 239 Anthyllis cytisoides L. (Leguminosae: Papilionoideae), 154 Antiaris africana Engl. (Moraceae): see il. toxicaría A. toxicaría ( R u m p h . ex Pers.) Leschen., 82, 83, 105, 172, 178, 181,186, 187 Antidesma venosum E . Meyer ex Tul. (Euphorbiaceae), 105, 189 Antrocaryon (Anacardiaceae), 74 A. micraster A . Chev. & Guillaumin, 81 Aphania senegalensis (Poir.) Radlk. (Sapindaceae), 129 Aphanocalyx (Leguminosae: Caesalpinioideae), 74 Aphloia (Flacourtiaceae), 237 A. theiformis (Vahl) Benn., 257, 258 Aphyllanthes (Liliaceae), 147 Apodocephala (Compositae), 232, 237 Apodytes dimidiata E . Meyer ex Arn. (Icacinaceae), 122, 165, 166,181,187,196,201,255,259 Apollonias barbujana (Cav.) Bornm. (Lauraceae), 246, 247, 249 Aporrhiza nítida Gilg ex Engl. (Sapindaceae), 91 Aptosimum (Scrophulariaceae), 193 A. depressum Burchell ex Benth.: see A. procumbens A. procumbens (Lehm.) Steud., 195 Aquilegia vulgaris L . (Ranunculaceae), 156 Arachis (Leguminosae, Papilionoideae), 178 Arbutus canariensis Veill. (Ericaceae), 246 A. pavarii Pampan., 159, 226 A. unedoL., 147,149, 152, 159 Archidium capense H o m s c h u c h (Archidiaceae), 182 Arctotis (Compositae), 141 Ardisiandra (Primulaceae), 162 Arenaria dyrís Humbert (Caryophyllaceae), 158 A. pungens Clemente ex Lagasca, 158 Argania (Sapotaceae), 226, 228, 229 A. spinosa (L.) Skeels, 54, 64, 149, 150, 154, 157, 225, 226, 227, 228, 229 Argemone mexicana L . (Papaveraceae), 254 Argyranthemum (Compositae), 246 Argyroderma (Aizoaceae), 140 Arístida (Gramineae), 140, 144, 193, 194, 210, 213, 221, 238, 239, 243 A. acutiflora Trin. & Rupr.: see Stipagrostis acut¡flora A. adoensis Hochst., 208 A. adscensionis L . , 114, 116, 120, 208, 209, 210, 212, 243, 252, 254, 267 A. barbicollis Trin. & Rupr., 114 A. cardosoi Cout., 252 A. 'coerulescens', 222 A. congesta R o e m . & Schult., 194, 195, 211, 242, 243 A. diffusa Trin., 141,195 A. funiculata Trin. & Rupr., 212, 213, 252 A. graciliflora Pilger: see A. stipitata A. hordeacea Kunth, 145 A. junciformis Trin. & Rupr., 194, 202
A. mutabilis Trin. & Rupr., 113, 116,120, 211, 212 A. pallida Steud.: see Arístida sieberana A. pungens Desf.: see Stipagrostis pungens A. rhiniochloa Hochst., 145, 208, 209, 210 A. rufescens Steud., 239, 242, 243 A. sieberana Trin., 176, 204, 207, 211, 212, 213 A. similis Steud., 239 A. stipitata Hackel, 101 A. stipoides L a m . , 206, 207 A. vanderystii D e Wild., 173 Artemisia (Compositae), 129 A. afra Willd., 129 A. campestris L., 229, 230 ss. glutinosa (J. G a y ) Battand., 222 A. gorgonum W e b b , 252 A. herba-alba Asso, 153, 222, 229, 230, 266 A. inculta Del.: see A. herba-alba A. tilhoana Quézel, 222 Arthraerua (Amaranthaceae), 137 A. leubnitziae (Kuntze) Schinz, 142,144 Arthraxon lancifolius Hochst. (Gramineae), 255 Arthrocarpum (Leguminosae: Papilionoideae), 111 Arthrocnemum (Chenopodiaceae), 267 A. dunense M o s s ex A d a m s o n , 144 A. glaucum (Del.) Ungern-Sternb., 224, 230 A. indicum (Willd.) M o q . , 144, 223, 263 Arthropteris orientalis (J. F . Gmelin) Posthumus (Oleandraceae), 99 Arundinaria (Gramineae), 237 A. alpina K . Schum., 55, 130, 167 A. marojejyensis A . C a m u s , 237 A. tesselata (Nees) M u n r o , 55,167 Arundo donax L . (Gramineae), 159 Ascarina (Chloranthaceae), 237 Ascarinopsis (Chloranthaceae), 232 A. coursii Humbert & Capuron, 237 Asclepias multicaulis Schltr. (Asclepiadaceae), 194 Ascolepis anthemiflora (Welw.) W e l w . (Cyperaceae), 100 A. elata W e l w . , 100 Aspalathus (Leguminosae: Papilionoideae), 132, 134 Asparagus (Liliaceae), 98, 156, 201 A. acutifolius L., 151, 152 A. albus L., 153,158 A. pastorianus W e b b & Berth., 229 A. stipularis Forssk., 158, 227, 266 A. warneckei (Engl.) Hutch., 83 Asphodelus (Liliaceae), 158, 221, 228 A. aestivus Brot., 152 A. fistulosus L . , 266 A. microcarpus Salzm. & Viv., 152, 158, 160 A. tenuifolius Cav., 229 Aspidium aculeatum Swartz (Aspidiaceae), 157 Aspilia mossambicensis (Oliver) Wild (Compositae), 115, 122 Asplenium (Aspleniaceae), 75 A. adiantum-nigrum L . , 157 A. dregeanum Kunze, 81 A. nidus L., 235 Aster (Compositae), 140 Asteriscus graveolens (Forssk.) D C . (Compositae), 221 Asteropeia densiflora Baker (Asteropeiaceae), 237 Asthenatherum forskalii (Vahl) Nevski (Gramineae), 144, 220 A. glaucum (Nees) Nevski, 191, 193 A. mossamedense (Rendle) Conert, 144 Astragalus (Leguminosae: Papilionoideae), 147 Asystasia gangetica (L.) T . Anderson (Acanthaceae), 182
Index ofplant names
Ataenidia (Marantaceae), 75 Atalaya (Sapindaceae), 199 A. natalensis R . A . Dyer, 199 Athyrium filix-femina (Lj Roth (Athyna.ceae), 151, 157 A triplex (Chenopodiaceae), 224, 267 A. halimus L . , 144, 223, 228, 229, 230, 266 A. mollis L., 230 A. vestita (Thunb.) Aellen, 267 Aubrevillea (Leguminosae: Mimosoideae), 74 A. kerstingii (Harms) Pellegr., 79 Aucoumea (Burseraceae), 74 A. klaineana Pierre, 77 A ugea (Zygophyllaceae), 137 Auxopus (Orchidaceae), 75 Avena bromoides (Gouan) Trabut (Grarnineae), 230 Avicennia (Avicenniaceae), 254, 261, 262, 263, 264 A. africana P. Beauv.: see A germinans A. germinans (L.) L., 261, 262 A. marina (Forssk.) Vierh., 253, 259, 261, 263, 264 A. nitida Jacq.: sec A. germinans Azanza garckeana (F. Hoffm.) Exell & Hillcoat (Malvaceae), 95,210 Azima tetracantha L a m . (Salvadoraceae), 182, 201 Azolla africana Desv. (Azollaceae), 265 Babiana (Iridaceae), 137, 141 Bachmannia (Capparidaceae), 199 Bafodeya benna (Scott Elliot) Prance (Chrysobalanaceae), 176 Baikiaea (Leguminosae: Caesalpinioideae), 90, 98 B. eminii Taubert: see B. insignis B. insignis Benth., 181 B. plurijuga H a r m s , 90, 97, 98,101 Baillonella (Sapotaceae), 74 Baissea wulfhorstii Schinz (Apocynaceae), 90 Balanites aegyptiaca (L.) Del. (Balanitaceae), 62, 87, 105, 107, 108, 121, 203, 206, 207, 209, 210, 212, 213, 214, 219, 222, 224 B. angolensis (Welw.) W e l w . ex Exell, 91, 95 B. maughamii Sprague, 90, 96, 200 B. orbicularis Sprague, 114, 121 B. wilsoniana D a w e & Sprague, 74,187,188 Ballochia (Acanthaceae), 111 Ballota (Labiatae), 228 B. hispánica (L.) M u n b y , 227 Balthasaria (Theaceae), 162 B. mannii (Oliver) Verde, 246, 253 B. schliebenii (Melchior) Verde, 246 Bambusa vulgaris Schrad. (Grarnineae), 55 Baphia burttii Baker f. (Leguminosae: Papilionoideae), 97 B. massaiensis Taubert, 90, 97 B. obovata Schinz: see B. massaiensis Barbeya (Barbeyaceae), 162 B. oleoides Schweinf., 115 Barleria (Acanthaceae), 140 B. hochstetteri Nees, 204 B. macrostegia Nees, 194 B. solitaria P . G . Meyer, 142 Barringtonia racemosa (L.) Sprengel (Lecythidaceae), 188, 261, 264 Barteria fistulosa Masters (Passifloraceae), 31 Bassia muricata (L.) Asch. (Chenopodiaceae), 230 Bathiaea (Leguminosae: Caesalpinioideae), 241 Bauhinia macrantha Oliver (Leguminosae: Caesalpinioideae): see B. petersiana B. natalensis Oliver, 201
329
B. petersiana C . Bolle, 90, 99 B. rufescens L a m . , 105, 206 B. taitensis Taubert, 114 B. tomentosa L . , 91 Beckeropsis uniseta (Nees) K . Schum. (Grarnineae): see Pennisetum unisetum Begonia (Begoniaceae), 165, 253 Beilschmiedia natalensis J. H . Ross (Lauraceae), 199 Bellevalia (Liliaceae), 158 Bequaertiodendron natalense (Sond.) Heine & J. H . Hemsley (Sapotaceae), 200 Berberís hispánica Boiss. & Reuter (Berberidaceae), 158 Berchemia discolor (Klotzsch) Hemsley (Rhamnaceae), 90, 95, 200, 201 B. zeyheri (Sond.) Grubov, 96 Berkheya (Compositae), 140, 194 B. onopordifolia ( D C . ) O . Hoffm. ex Burtt Davy, 194 B. rígida (Thunb.) Bolus & Wolley D o d ex A d a m s o n & T . M . Salter, 194 Berkheyopsis angolensis O . Hoffm. (Compositae), 144 Berlinia auriculata Benth. (Leguminosae: Caesalpinioideae), 83 B. giorgii D e Wild., 90, 173 B. grandiflora (Vahl) Hutch. & Dalz., 176 B. occidentalis Keay, 77 Berzelia lanuginosa Brongn. (Bruniaceae), 134 Betula alba auct.: see B. péndula, Betulaceae B. péndula Roth, 147, 149,156 Bidens pilosa L. (Compositae), 252 Biscutella (Cruciferae), 228 Bivinia (Flacourtiaceae), 186 B.jalbertii Tul., 188 Blaeria (Ericaceae), 132,168 B. mannii (Engl.) Engl., 74 B. spicata Hochst. ex A . Rich., 211 Blechnum spicant (L.) Roth (Blechnaceae), 157 Blepharis (Acanthaceae), 140 B. acanthoides sensu D . B . Burtt, 116 B. ciliaris (L.) B . L. Burtt, 108, 204 B. edulis (Forssk.) Pers.: see B. ciliaris B. linariifolia Pers., 120, 212 B. maderaspatensis (L.) Roth, 90 Blighia unijugata Baker (Sapindaceae), 199 Boerhavia coccínea Miller (Nyctaginaceae), 206 B. repens L., 252 Bolbitis (Lomariopsidaceae), 75 Bolusanthus (Leguminosae: Papilionoideae), 87 B. speciosus (Bolus) H a r m s , 96 Bombax costatum Pellegr. & Vuillet (Bombacaceae), 105,109 Bonamia poranoides Hallier f. (Convolvulaceae), 182 Borassus aethiopum Martius (Palmaceae), 83, 85, 95, 105, 107, 108,189,201,209 B. madagascariensis Bojer, 243 Boscia (Capparidaceae), 111 B. albitrunca (Burchell) Gilg & C . Benedict, 90, 137, 140, 191,193,194 B. angustifolia A . Rich., 87, 207 B. coriácea Pax, 114, 119 B. foetida Schinz, 140 B. microphylla Oliv., 95 B. rehmanniana Pest, 95 B. salicifolia Oliver, 87,105, 109, 210, 222 B. senegalensis (Pers.) L a m . ex Poir., 206, 207, 210, 212, 213,216 Bosqueia angolensis Ficalho: see Trilepisium madagascariense B. phoberos: see T. madagascariense
330
Index ofplant names
Boswellia (Burseraceae), 111 5. ameero Balf.f., 115 B. dalzielii Hutch., 105, 107 B. elongata Balf.f., 115 B. hildenbrandtii Engl.: see B. neglecta B. neglecta S. Moore, 114, 120 B. papyrifera (Del.) Hochst., 107, 203, 209, 210, 211 B. socotrana Balf.f., 115 Bothriochloa insculpta (Hochst.) A . C a m u s : see Dichanthium insculptum Bottegoa (Sapindaceae), 111 Bowringia mildbraedii H a r m s (Leguminosae: Papilionoideae), 81 Brabeium stellatifolium L . (Proteaceae), 135 Brachiaria (Gramineae), 169 B. brizantha (Hochst. ex A . Rich.) Stapf, 85, 101, 173, 174, 210 B. eruciformis (Smith) Griseb., 114 B. falcifera (Trin.) Stapf, 178 B. fulva Stapf: see B. jubata B.jubata (Fig. & D e Not.) Stapf, 107 B. lata (Schumach.) C . E . Hubbard, 209 B. leersioides (Hochst.) Stapf, 114 B. mutica (Forssk.) Stapf, 266 B. nana Stapf, 243 B. nigropedata (Munro ex Ficalho & Hiern) Stapf, 193 B. ramosa L . , 243 B. serrata (Thunb.) Stapf, 194, 196 Brachylaena (Compositae), 236 B. discolor D C , 166 B. huillensis O . Hoffm., 117,188 B. hutchinsii Hutch.: see B. huillensis B. ilicifolia (Lam.) E . P. Phillips & Schweick., 201 B. microphylla Humbert, 237 B. uniflora Harv., 199 Brachypodium perrieri A . C a m u s (Gramineae), 239 B. ramosum (L.) R o e m . & Schult., 154 Brachystegia (Leguminosae: Caesalpinioideae), 87, 92, 96, 97, 106 B. allenii Burtt Davy & Hutch, 92, 93 B. angustistipulata D e Wild., 92 B. bakerana Burtt Davy & Hutch., 63, 89, 92, 98 B. boehmii Taubert, 30, 54, 92, 93, 99 B. bussei H a r m s , 92, 93 B. cynometroides Harms, 77 B. floribunda Benth., 93, 99 B. glaberrima R . E . Fries, 92, 93 B. glaucescens Burtt Davy & Hutch.: see B. microphylla B. laurentii (De Wild.) Louis ex Hoyle, 77, 78 B. leonensis Burtt Davy & Hutch., 77 B. longifolia Benth., 91, 92, 93, 97 B. manga D e Wild., 92, 93 B. microphylla Harms, 92, 93, 99 B. mildbraedii Harms, 77 B. puberula Burtt Davy & Hutch., 92, 97 B. russelliae I. M . Johnston, 92 B. spiciformis Benth., 54, 91, 92, 93, 97, 98, 99, 173, 188 B. stipulata D e Wild., 92, 99 B. tamarindoides W e l w . ex Benth., 92, 93 B. taxifolia Harms, 91, 92, 93, 99 B. torrei Hoyle, 92 B. utilis Burtt Davy & Hutch., 92, 93 B. wangermeeana D e Wild., 92, 93, 97, 173 Brackenridgea arenaria (De Wild. & T h . Durand) N . Robson (Ochnaceae), 173
Brenania (Rubiaceae), 74 Breonadia microcephala (Del.) Ridsdale: seeAdina microcephala Breonia sp. (Rubiaceae), 255 Bridelia ferruginea Benth. (Euphorbiaceae), 85,105, 174 B. taitensis Pax, 114 Bromus erectus Hudson (Gramineae), 157 B. madritensis L . , 228 B. rubens L . , 228 B. speciosus Nees, 169 Broussonetia greveana (Bâillon) C . C . Berg (Moraceae), 241 Brucea antidisentérica Miller (Simaroubaceae), 122 Bruguiera (Rhizophoraceae), 261 B. gymnorrhiza (L.) L a m . , 259, 261, 263, 264 Brunia (Bruniaceae), 134 Bryonia dioica Jacq. (Cucurbitaceae), 227 Bryum argenteum Hedwig (Bryaceae), 182 Buchholzia (Capparidaceae), 74 Buchnerodendron speciosum Giirke (Flacourtiaceae), 81 Buddleja corrugata (Benth.) E . P. Phillips (Loganiaceae), 195 B. saligna Willd., 193, 194, 195 B. salviifolia L a m . , 195 Bulbine (Liliaceae), 141 Bulbophyllum (Orchidaceae), 236, 237 B. leptostachysum Schltr., 238 Bulbostylis abortiva (Steud.) C . B . Clarke (Cyperaceae), 84 B. basalis Fosberg, 259 B. cinnamomea C . B . Clarke, 100 B. firingalavensis Chermezon, 243 B. laniceps C . B . Clarke ex T h . Durand & Schinz, 84 B. xerophila Chermezon, 243 Bupleurum spinosum G o u a n (Umbelliferae), 156,158 Burchellia (Rubiaceae), 199 Burkea africana H o o k . (Leguminosae: Caesalpinioideae), 85, 87, 95, 96, 97,99,105,106, 107,173,196 Burmannia (Burmanniaceae), 75, 84 Burttdavya nyasica Hoyle (Rubiaceae), 97,186 Burttia prunoides Baker f. & Exell (Connaraceae), 97 Bussea massaiensis (Taubert) H a r m s (Leguminosae: Caesalpinioideae), 97 Butyrospermum (Sapotaceae), 103 B. paradoxum (Gaertner f.) Hepper, 55, 83, 85, 105, 107, 108 B. parkii (G. D o n ) Kotschy: see B. paradoxum Buxus baleárica L a m . (Buxaceae), 153 B. hildebrandtii Bâillon, 115 B. sempervirens L . , 156 Byrsocarpus orientalis (Bâillon) Baker (Connaraceae), 90, 98 Cadaba(Capparidaceae), 111,242 C. aphylla (Thunb.) Wild, 96, 201 C. farinosa Forssk., 114, 206 C. glandulosa Forssk., 113, 204, 212, 213 C. heterotricha Hook., 114 Cadia purpurea (Picciv.) Aiton (Leguminosae: Papilionoideae), 115 Caesalpinia trothae H a r m s (Leguminosae: Caesalpinioideae), 114 Cajanus cajan (Leguminosae: Papilionoideae), 252 Calamus (Palmaceae), 83 Calendula algeriensis Boiss. & Reuter (Compositae), 158, 227 C. murbeckii Lanza, 229 Calicotome intermedia C . Presl. (Leguminosae: Papilionoideae): see C. villosa C. villosa (Poir.) Link, 152, 158, 226 Calligonum (Polygonaceae), 220
Index ofplant names
C. comosum L'Hér., 204, 207, 219, 224 Calluna vulgaris (L.) Hull (Ericaceae), 147, 152, 247 Calodendrum capense (L.f.) Thunb. (Rutaceae), 115, 130, 166, 196,199 Caloncoba glauca (P. Beauv.) Gilg (Flacourtiaceae), 81 C. welwitschii (Oliver) Gilg, 80 Calophyllum eputamen P. F . Stevens (Guttiferae), 258 C. inophyllum L., 257 C. tacamahaca Willd., 258 Calotropis procera (Aitón) Aitón f. (Asclepiadaceae), 121, 212, 219,251,252 Calpocalyx (Leguminosae: Mimosoideae), 74 Calvaría galeata A. W . Hill: see Sideroxylon galeatum C. major Gaertner f.: see S. majus Calyptrotheca (Portulacaceae), 111 C. somalensis Gilg, 114 C. taitensis (Pax & Vatke) Brenan, 114 Campnosperma seychellarum Marchand (Anacardiaceae), 257 Campylanthus salsoloides (L.f.) Roth (Scrophulariaceae), 252 Canarina abyssinica Engl. (Campanulaceae), 246 C. canariensis (L.) Vatke, 246 C. eminii Asch. ex Schweinf., 246 Canarium (Burseraceae), 235,236 C. mauritianum Blume: see C. paniculatum C. paniculatum (Lam.) Benth. ex Engl., 258 C. schweinfurthii Engl., 78, 79, 80, 81, 82, 91,172,173,181 Canavalia rosea (Swartz) D C . (Leguminosae: Papilionoideae), 253 Canthium (Rubiaceae), 83, 98 C. bibracteatum (Baker) Hiern, 259 C. burttii Bullock, 98, 99 C. frángula S. Moore, 90 C. keniense Bullock, 115 C. lactescens Hiern, 99 C. martinii Dunkley, 90 C. schimperanum A . Rich., 182 C. vulgare (K. Schum.) Bullock, 182 Capitanya (Labiatae), 111 Capparis (Capparidaceae), 189,219 C. decidua (Forssk.) Edgew., 219,224 C. elaeagnoides Gilg: see C. fascicularis C. erythrocarpos Isert, 91, 129,178 C. fascicularis DC, 115, 176,182 C. sepiaria L . , 201 C. tomentosa L a m . , 182,183 Caperonia palustris (L.) A . St. Hil. (Euphorbiaceae), 108 Caralluma (Asclepiadaceae), 114, 116,139 C. edithae N . E . Br., 116 C. penicillata (Defl.) N . E . Br., 116 Carapa grandiflora Sprague (Meliaceae), 85, 181 C. procera D C , 82, 83 Cardamine (Cruciferae), 236 Carex capillaris L . (Cyperaceae), 147, 158 C.distachya Desf., 152 C. distans L . , 156 C. leporina L., 155 Carica papaya L . (Caricaceae), 252 Carissa (Apocynaceae), 188, 189 C. bispinosa (L.) Desf. ex Brenan, 96, 135, 201 C. edulis Vahl, 98,109, 115, 121, 129,182 C. haematocarpa (Eckl.) A . D C , 137, 201 C. xylopicron Thouars, 258 Carphalea glaucescens (Klotzsch) Verde. (Rubiaceae), 114 Carpodiptera africana Masters (Tiliaceae), 189 Carthamnus fruticosus Maire (Compositae), 153
331
Carum verticillatum (L.) Koch (Umbelliferae), 155 Casearia barteri Masters (Flacourtiaceae), 211 C. battiscombei R . E . Fries, 122 C. gladiiformis Masters, 200 Cassia (Leguminosae: Caesalpinioideae), 98, 241 C. abbreviata Oliver, 95, 96, 99' ss. kassneri (Baker) Brenan, 114 C. aschrek Forssk.: see C. itálica C. itálica (Miller) L a m . ex F . W . Andrews, 219 C. mimosoides L., 178 C. sieberana D C , 105 C singueana Del., 189 C. tora L., 212 Cassine (Celastraceae), 98 C aethiopica Thunb., 91, 182, 201, 243, 259 C. buchananii Loes., 115,129 C. parvifolia Sond., 134 C. peragua L., 135 Cassinopsis ilicifolia (Hochst.) Kuntze (Icacinaceae), 195 Cassipourea (Rhizophoraceae), 187 C. annobonensis Mildbr., 253 C. congoensis R . Br. ex D C , 83, 122, 166 C. euryoides Alston, 188 C. gerrardii (Schinz) Alston, 200 C. gossweileri Exell, 98 C. gummiflua Tul., 253 C. malosana (Baker) Alston: see C. congoensis Casuarina equisetifolia L. (Casuarinaceae), 259 Catophractes alexandri D . D o n (Bignoniaceae), 95 Caucanthus albidus (Niedenzu) Niedenzu (Malphigiaceae), 114 Cavacoa quintasii (Pax & Hoffm.) J. Léonard (Euphorbiaceae), 253 Caylusea canescens (Murray) W e b b (Resedaceae): see C. hexagyna C. hexagyna (Forssk.) M . L . Green, 219, 220 Cedrus atlántica (Endl.) Carrière (Pinaceae), 60, 146, 147, 149, 150,151, 152, 154, 155, 156,157 C . brevifolia (Hook.f.) A . Henry, 155 C. deodara Loudon, 155 C . libani A . Rich., 155 Ceiba pentandra (L.) Gaertner (Bombacaceae), 81, 83, 105, 178, 253 Celsia insularis M u r b . : see Verbascum capitis-viridis Celtis (Ulmaceae), 181 C. africana Burm.f., 80, 191,194, 195, 196, 200 C. australis L., 149 C. brownii Rendle, 80, 83 C. durandii Engl.: see C . gomphophylla C. gomphophylla Baker, 90,199, 253 C. integrifolia L a m . , 105, 209 C. mildbraedii Engl., 79, 178, 199, 253 C. philippensis Blanco: see C. brownii C. prantlii Priemer ex Engl., 253 C. wightii Planchón, 187 C. zenkeri Engl., 79, 80, 172 Cenchrus biflorus Roxb. (Gramineae), 206, 207, 210, 212, 216 C.ciliarisL., 114,116 C. prieurii (Kunth) Maire, 210 Centaurea (Compositae), 147 Centauropsis (Compositae), 232, 237 Cephaelis peduncularis Salisb.: see Psychotria peduncularis Cephalocroton socotranus Balf.f. (Euphorbiaceae), 115 Cephalopentandra (Cucurbitaceae), 111 Cephalosphaera (Myristicaceae), 186 C. usambarensis (Warb.) W a r b . , 187
332
Index ofplant names
Ceraria (Portulacaceae), 137 C. longepedunculata M e r x m . & Podl, 140 C. namaquensis (Sond.) H . Pearson, 140 Cerastium (Caryophyllaceae), 236 Ceratonia (Leguminosae: Caesalpinioideae), 147, 226 C. siliqua L . , 149, 152,158,159, 226, 228, 252 Ceratophyllum demersum L . (Ceratophyllaceae), 265 Ceriops tagal (Pers.) C . B . Robinson (Rhizophoraceae), 259, 261,263,264 Ceropegia (Asclepiadaceae), 111 C. dimorpha Humbert, 238 Chaetacme aristata Planchón (Ulmaceae), 129, 166,199, 253 Chaetocarpus africanus Pax (Euphorbiaceae), 80 Chamaemeles (Rosaceae), 247 Chamaerops (Palmaceae), 147 C. humilis L . , 149, 152, 157,158, 160, 228 Chascanum marrubifolium Fenzl ex W a l p . (Verbenaceae), 204 Cheilanthes (Sinopteridaceae), 239 Chenolea tomentosa (Lowe) Maire (Chenopodiaceae), 229 Chenopodium (Chenopodiaceae), 122 C. ambrosioides L . , 145 Chidlowia (Leguminosae: Caesalpinioideae), 74 Chionanthus foveolatus (E. Meyer) Steam (Oleaceae), 135, 165 Chionothrix (Amaranthaceae), 111 Chloris gay ana Kunth (Gramineae), 126, 208 C . prieurii Kunth, 176 •C. roxburghiana Schultes, 114, 116 C. virgata Swartz, 194, 212, 213, 243 Chlorophora (Moraceae), 187 C. excelsa (Welw.) Benth., 74, 79, 80, 81, 172, 181, 186, 187,188,189,253 C. greveana (Bâillon) Leandri: see Broussonetia greveana C. regia A . Chev., 178 Chondropetalum mucronatum (Masters) Pillans (Restionaceae), 134 Chrozophora brocchiana Vis. (Euphorbiaceae), 204, 219 Chrysalidocarpus (Palmaceae), 232, 237 C. acuminum Jumelle, 238 C. decipiens Becc, 237 Chrysanthemoides monilifera (L.) Norlindh (Compositae), 135 Chrysanthemum (Compositae), 158 Chrysithrix (Cyperaceae), 132 Chrysocoma (Compositae), 135,137, 139, 140 C tenuifolia Bergius, 193, 194, 195, 196 Chrysophyllum albidum G . D o n (Sapotaceae), 181, 253 C . boivinianum (Pierre) J. H . Hemsley, 255 C. gorungosanum Engl., 164, 181 C. perpulchrum Mildbr. ex Hutch. & Dalz., 74, 79, 82, 187 C . viridifolium W o o d & Franks, 167, 199 Chrysopogon aucheri (Boiss.) Stapf (Gramineae): see C. plumulosus C. plumulosus Hochst., 116, 121, 222 Cicca disticha L . (Leguminosae: Papilionoideae), 252 Cincinnobotrys (Melastomataceae), 162 Cinnamomum zeylanicum Nees (Lauraceae), 257 Cissus (Vitaceae), 242 C. cactiformis Gilg, 128 C. petiolata Hook.f., 182 C. quadrangularis L . , 81, 114, 117, 128, 182, 201 C. rotundifolia (Forssk.) Vahl, 114,182 Cistanche phelipaea (L.) Cout. (Orobanchaceae), 229 Cistus (Cistaceae), 147, 226, 228 C . clusii Dunal, 154 C. crispusL., 152 C. laurifolius L . , 157
C. parviflorus L a m . , 159 C. populifolius L . , 152 C. sahiifolius L . , 152 C. symphytifolius L a m . , 249 C villosusL., 154 Citropsis daweana Swingle & M . Kellerman (Rutaceae), 90 Citrullus colocynthis (L.) Schrader (Cucurbitaceae), 219 C. ecirrhosus Cogn., 144 Citrus (Rutaceae), 159 Cladium mariscus (L.) Pohl (Cyperaceae), 265 Cladonia (Cladoniaceae), 237 C . medusina (Bory) Nylander, 189 C. pycnoclada (Persoon), Nylander, 239 Cladostigma (Convolvulaceae), 111 Clausena anisata (Willd.) Hook.f. ex Benth (Rutaceae), 81, 122, 173 Cleistanthus polystachyus H o o k , ex Planchón (Euphorbiaceae), 187 C . schlechteri (Pax) Hutch., 200 Cleistochlamys (Annonaceae), 87 Clematis cirrhosa L . (Ranunculaceae), 151, 152,153, 154, 227 C. flammula L . , 158 Cleome (Capparidaceae), 144 C. scaposa D C , 204 C . viscosa L . , 252 Clerodendrum (Verbenaceae), 237 C . glabrum E . Meyer, 200 Clethra arbórea Aitón (Clethraceae), 246, 247 Cliffortia (Rosaceae), 132, 134 C . arbórea Marloth, 134 C . grandifolia Eckl. & Zeyh., 134 Cocos (Palmaceae), 189 C . nucífera L . , 257, 259 Coelocaryon (Myristicaceae), 74 C . botryoides Vermoesen, 83 Coffea arabica L . (Rubiaceae), 252 Cola clavata Masters (Sterculiaceae), 187 C. cordifolia (Cav.) R . Br., 178 C. digitata Masters, 253 C . gigantea A. Chev., 79 C. greenwayi Brenan, 164 C. laurifolia Masters, 176 C. natalensis Oliver, 199 Colchicum (Liliaceae), 158 Colea seychellarum Seem., (Bignoniaceae), 257 Coleochloa setífera (Ridley) Gilly (Cyperaceae), 238 Colocasia antiquorum Schott (Araceae): see C. esculenta C. esculenta (L.) Schott, 252 Colocynthis vulgaris Schrader: see Citrullus colocynthis Colophospermum (Leguminosae: Caesalpinioideae), 87 C. mopane (Kirk ex Benth.) J. Léonard, 31, 54, 61, 62, 89, 94,140, 141, 143, 144,191 Combretodendon africanum (Welw. ex Benth.) Exell: see Petersianthus macrocarpum C. macrocarpum P. Beauv.: see P. macrocarpum Combretum (Combretaceae), 75, 98, 110, 129, 173, 241 C. aculeatum Vent., 113, 114, 141, 209, 212 C . apiculatum Sond., 95, 96, 193, 201 C . camporum Engl., 90, 172 C. celastroides W e l w . ex Lawson, 90, 97, 98 ss. laxiflorum (Welw. ex Lawson) Exell, 173 ss. orientale Exell, 97 C . collinum Fresen., 85, 87, 90, 95, 96, 97, 98, 105, 107, 109, 189,193,201,209,210 C. cordofanum Engl. & Diels: see C. glutinosum
Index of plant names
C. C. C. C. C.
elaeagnoides Klotzsch, 90 erythrophyllum (Sond.) Burchell, 141 fragrans F . Hoffm., 95, 105 ghasalense Engl. & Diels: see C. fragrans glutinosum D C , 105, 106, 107, 108, 203, 209, 210, 212, 213 C. hartmannianum Schweinf., 62, 107 C. hereroense Schinz, 96 C. imberbe W a w r a , 91, 95, 96, 141, 201 C. kraussii Hochst., 165, 199 C. mechowianum O . Hoffm.: see C. collinum C. micranthum G . D o n , 109 C. molle R . Br. ex G . D o n , 87, 95, 96, 105, 121, 129, 196, 201,211 C. mossambicense (Klotzsch) Engl., 90 C. mucronatum Schumach. & Thonn., 81 C. nigricans Lepr. ex Guill. & Perr., 105, 107 C. oxystachyum W e l w . ex Lawson, 95 C. paniculatum Vent., 81, 209 C. psidioides W e l w . , 97,173 C. racemosum P . Beauv., 81 C. schumannii Engl., 187, 188 C. trothae Engl. & Diels: see C . celastroides ss. orientale C. zeyheri Sond., 96, 97, 201 Commelina benghalensis L . (Commelinaceae), 191 Commicarpus (Nyctaginaceae), 111 C. verticillatus (Poir.) Standley, 252 Commidendrum (Compositae), 254 C. robustum D C , 254 C. rugusum (Aitón) D C , 254 C. spurium D C , 254 Commiphora (Burseraceae), 31, 63, 98, 110, 111, 113, 114, 115, 116, 117, 120, 121, 125, 128, 140, 144, 145, 181, 188, 189,191 C. africana (A. Rich.) Engl., 87, 105, 107, 114, 193, 203, 206, 207, 208, 209, 210, 211, 212, 216 C. anacardiifolia Dinter & Engl., 95 C. angolensis Engl., 90, 95,193 C. baluensis Engl., 116 C. boiviniana Engl., 114 C. campes tris Engl., 114, 116 C. capensis (Sond.) Engl., 140 C. dalzielii Hutch., 176 C. dulcis Engl.: see C saxícola C. engleri Guillaumin, 116 C. erythraea (Ehrenb.) Engl., 114 C. gracilifrondosa Dinter ex V a n der Walt, 140 C harveyi (Engl.) Engl., 196, 199, 200 C. madagascariensis Jacq., 128 C. merkeri Engl., 116, 128 C. mollis (Oliver) Engl, 96, 114 C. monstruosa ( H . Perrier) Capuron, 242 C. mossambicensis (Oliver) Engl., 99 C. namaensis Schinz, 140 C. oblanceolata Schinz, 140 C. pedunculata (Kotschy & Peyr.) Engl., 105 C. pyracanthoides Engl., 95, 96 C. riparia Engl.: see C. mollis C. saxícola Engl., 140 C. schimperi (Berger) Engl., 114,116, 128 C. trothae Engl.: see C schimperi Conocarpus erectus L . (Combretaceae), 253, 261, 262 Convolvulus gharbensis Battand. & Pitard (Convolvulaceae), 158 C. trabutianus Schweinf. & Muschler, 229 C. tricolor L . , 158
333
Conyza (Compositae), 237 C . pinnata (L.f.) Kuntze, 194 Corchorus (Tiliaceae), 252 C. tridens L . , 206 Cordeauxia (Leguminosae: Caesalpinioideae), 111 Cordia abyssinica R . Br. (Boraginaceae), 209, 210, 211 C . caffra Sond., 199, 200 C. gharaf{¥brssk.) Ehrenb. ex Asch.: see C. sinensis C. millenii Baker, 181 C. ovalis R . Br. ex D C , 114, 128, 129,182 C. rothii R o e m . & Schult.: see C. sinensis C. sinensis L a m . , 114, 128, 144, 206, 212, 222 C . subcordata L a m . , 257 Cordyla africana Lour. (Leguminosae: Caesalpinioideae), 96, 186, 187 C. madagascariensis R . Viguier, 241 Coriandrum (Umbelliferae), 147 Cornulaca monacantha Del. (Chenopodiaceae), 204, 216, 220, 221,224 Coronilla glauca L . (Leguminosae: Papilionoideae): see C . valentina C. valentina L . , 151 Cosmos (Compositae), 243 Costus (Zingiberaceae), 75 Cotoneaster fontanesii Spach (Rosaceae), 151 Cotula (Compositae), 141 C. coronopifolia L . , 145 Cotyledon (Crassulaceae), 137, 140 C . decussata Sims, 195 C. orbiculata L . , 142 C. paniculata L . , 140 Coula (Olacaceae), 74 C . edulis Bâillon, 77 Crabbea acaulis N . E . Br. (Acanthaceae), 194 Craibia brevicaudata (Vatke) D u n n (Leguminosae: Papilionoideae) ss. burtii (Baker f.) J. B . Gillett, 97 C. zimmermannii (Harms) H a r m s ex D u n n , 200 Craspedorhachis africana Benth. (Gramineae), 239 Crassula (Crassulaceae), 115,132, 137, 140,201 C. arborescens (Miller) Willd., 140 C . portulacea L a m . , 137, 201 Crataegus (Rosaceae), 156 C . azarolus L . , 149 C . laciniata Ucria, 156 C monogynal&cq., 149, 151,157,158 Craterispermum laurinum (Poir.) Benth. s.l. (Rubiaceae), 82, 91 C. montanum Hiern, 253 Crocus (Iridaceae), 158 C . boulosii Greuter, 226 Crossandra nilotica Oliver (Acanthaceae), 176 Crossopteryx febrífuga (Afzel. ex G . D o n ) Benth. (Rubiaceae), 83, 85, 105, 189 Crotalaria(Leguminosae: Papilionoideae), 111, 140, 243 C . agatiflora Schweinf. ss. imperialis (Taub.) Polhill, 130 C. microphylla Vahl, 204 C.podocarpaBC., 191 C . retusa L . , 252 Crotón (Euphorbiaceae), 236 C dichogamus Pax, 115, 128, 129, 182 C . dybowskii Hutch., 172 C . gratissimus Burchell, 90,193, 200 C . macrostachyus Hochst. ex Del., 130 C . megalobotrys Muell. Arg., 91, 95
334
Index ofplant names
Crotón (Euphorbiaceae)—contd C. megalocarpus Hutch., 167, 181 C. mubango Muell. Arg., 81 C. pseudopulchellus Pax, 90, 188 C. scheffleri Pax, 90 C. socotranus Balf.f., 255 C. stelluliferus Hutch., 253 C. sylvaticus Hochst., 74,199 Crotonogyne (Euphorbiaceae), 74 Crudia gabonensis Pierre ex H a r m s (Leguminosae: Caesalpinioideae), 76 Cryptocarya angustifolia E . Meyer ex Meissner (Lauraceae), 134 C latifolia Sond., 165 C. woodii Engl., 165 Cryptosepalum (Leguminosae: Caesalpinioideae), 63 C. pseudotaxus Baker f., 57, 90, 97 C. staudtii H a r m s , 77 C. tetraphyllum (Hook.f.) Benth., 82 Ctenium concinnum Nees (Gramineae), 194, 239 C. elegans Kunth, 209 C. newtonii Hack., 85, 173, 210 C. somátense (Chiov.) Chiov., 210 Cunonia capensis L . (Cunoniaceae), 135 Cuphocarpus (Araliaceae), 232, 236 Cupressus atlántica Gaussen (Cupressaceae), 146, 149, 150, 152, 153 C. dupreziana A . C a m u s , 47, 218, 222 C. sempervirens L., 146,149, 150,152, 153,159, 226 Curtisia (Cornaceae), 162 C. dentata (Burm.f.) C . A . Smith, 165 C. fa*ginea Aitón: see C. dentata Cussonia (Araliaceae), 201, 237 C. arbórea Hochst. ex A . Rich., 83, 85,105 C. barteri Seemann: see C. arbórea C.holstii Engl., 115 C kirkii Seemann: see C. arbórea C. paniculata Eckl. & Zeyh., 194, 195 C. sessilis Lebrun, 173 C. spicata Thunb., 135, 194 C. zimmermannii Harms, 117, 188, 189 Cyanotis nodiflora Kunth (Commelinaceae), 238 Cyathea (Cyatheaceae), 165 C. manniana Hook., 82 Cyclamen (Primulaceae), 147 C. rohlfsianum Asch., 226 Cyclopia (Leguminosae: Papilionoideae), 132 Cydonia oblonga Miller (Rosaceae), 252 Cylicodiscus (Leguminosae: Mimosoideae), 74 Cylicomorpha parviflora Urban (Caricaceae), 164, 187 Cymbopogon (Gramineae), 50, 193, 210 C. commutatns (Steudel) Stapf, 208 C. excavatus (Hochst.) Stapf ex Burtt Davy, 202, 210 C. giganteas (Hochst.) Chiov., 210 C. nervatus Chiov., 108, 212, 213 C. plicatus Stapf, 239 C. plurinodis (Stapf) Stapf ex Burtt Davy, 137, 194 C. pospischilii (K. Schum.) C . E . Hubbard: see C. plurinodis C. proximus (Hochst. ex A . Rich.) Stapf: see C schoenanthus C. schoenanthus (L.) Sprengel, 208, 209, 211,212, 221 C. validus Stapf ex Burtt Davy, 201 Cynodon dactylon (L.) Pers. (Gramineae), 50, 116, 126, 127, 128, 194, 210, 246, 253 C. hirsutus Stent, 195 C. incompletus Nees, 194
Cynometra (Leguminosae: Caesalpinioideae), 235 C. alexandri C . H . Wright, 78, 85,172, 181, 182 C. ananta Hutch. & Dalz., 77 C. hankei H a r m s , 77 C. leonensis Hutch. & Dalz., 77 C. mannii Oliver, 253 C. megalophyllaHaims, 178 C. vogelii Hook.f., 176 C. webberi Baker f., 188 Cynomorium coccineum L. (Cynomoriaceae), 229 Cynorkis (Orchidaceae), 238 Cynosurus echinatus L. (Gramineae), 157 Cyperus esculentus L . (Cyperaceae), 100 C. haspan L . , 265, 266 C. laevigatus L., 144, 145, 223, 267 C. margaritaceus Vahl, 100 C. obtusiflorus Vahl, 194 C. papyrus L . , 55, 264, 265, 266 C. platycoulis Baker, 100 Cyphostemma currorii (Hook.f.) Descoings (Vitaceae), 140 Cyrtosperma senegalense (Schott) Engl. (Araceae), 83, 266 Cytisus albidus D C . (Leguminosae: Papilionoideae), 227 C. arbóreas (Desf.) D C , 151 C. balansae Ball, 158 C. battandieri Maire, 151, 156, 157 C. linifolius L a m . , 151 C. maurus Humbert & Maire, 152 C. monspessulanus L., 152 C. proliferus L.f., 248 C. stenopetalus (Webb & Berth.) Christ, 246, 251, 252 C. triflorus L'Hérit.: see C. villosus C. villosus Pourret, 152, 157 Daboecia azorica Tutin & E . W a r b . (Ericaceae), 247 Dacryodes edulis (G. D o n ) H . J. L a m (Burseraceae), 91,172, 253 Dactylis glomerata L . (Gramineae), 152 Dactyloctenium (Gramineae), 267 D. aegyptium (L.) Willd., 210 D. geminatum Hack., 263 D. giganteum Fisher & Schweick., 101 D. pilosum Stapf, 259 D. robecchii Chiov., 116 D. sp., 126 Dalbergia (Leguminosae: Papilionoideae), 235, 236, 241 D. armata E . Meyer, 96, 200, 201 D. boehmii Taubert, 95 D. ecastaphyllum (L.) Taubert, 253 D. hostilis Benth., 109 D. martinii F . White, 90 D. melanoxylon Guill. & Perr., 96, 105, 107, 116, 189, 208, 209,210,212,213 D. obovata E . Meyer, 196,200 Daniellia alsteeniana Duvign. (Leguminosae: Caesalpinioideae), 90,173 D. ogea (Harms) Rolfe ex Holland, 178 D. oliveri (Rolfe) Hutch. & Dalz., 83, 85,105, 106,107 Danthonia (Gramineae), 134 D. forskalii (Vahl) R . Br.: see Asthenatherum forskalii D. macowanii Stapf: see Merxmuellera macowanii D. mossamedensis Rendle: see A mossamedensis Danthoniopsis dinteri (Pilger) C . E . Hubbard (Gramineae), 145 Daphne gnidium L . (Thymelaeaceae), 151, 157, 158, 249 D. laureola h., 151,156 Dasysphaera (Amaranthaceae), 111 D. prostrata (Gilg) Cavaco, 120
Index of plant names
Daucus (Umbelliferae), 221 Decaryia madagascariensis Choux (Didiereaceae), 242 Deckenia (Palmaceae), 257 D. nobilis (Moore) H . A . Wendl. ex Balf.f., 257 Decorsella (Violaceae), 74 Delonix adansonioides (R. Viguier) Capuron (Leguminosae: Caesalpinioideae), 242 D. elata (L.) Gamble, 114 D. regia (Bojer) Rafin., 241 Dendrosicyos (Cucurbitaceae), 111 D. socotranus Balf.f., 255 Desbordesia (Irvingiaceae), 74 Deschampsia (Gramineae), 169 Desmanthus virgatus Willd. (Leguminosae: Mimosoideae), 252 Desmodium torttiosum (Swartz) D C . (Leguminosae: Papilionoideae), 252 Detarium microcarpum Guill. & Perr. (Leguminosae: Caesalpinioideae), 31, 105,107 D. senegalense J. F. Gmelin, 85,178 Dialium engleranum Henriques (Leguminosae: Caesalpinioideae), 85, 97,173 D. guineenseWm., 105, 178, 253 D. schlechteri Harms, 200 Dichanthium insculptwn (A. Rich.) W . D . Clayton (Gramineae), 121,128 Dichapetalum (Dichapetalaceae), 241 Dichrostachys (Leguminosae: Mimosoideae), 242 D. cinérea (L.) Wight & Arn., 84, 85, 96, 105, 107, 189, 193, 207,208, 209,210,212,214,251 Dicksonia arborescens L'Hérit. (Dicksoniaceae), 254 Dicoma carbonaria Humbert (Compositae), 242 D. foliosa O . Hoffm., 145 D. incana (Baker) O . Hoffm., 237, 242, 243 D. macrocephala D C , 194 D. oleifolia Humbert, 243 Dicoryphe (Hamamelidaceae), 232 D. viticoides Baker, 237 Dicraeopetalum (Leguminosae: Papilionoideae), 111 Dicranopteris linearis (Burm.) Underw. (Gleicheniaceae), 257 Dictyosperma album (Bory) H . A . Wendl. (Palmaceae), 257, 258 D. aurewn Balf.f.: see D. album Didelotia (Leguminosae: Caesalpinioideae), 74 D. brevipaniculata J. Léonard, 77 D. idae Oldeman, de Wit & J. Léonard, 76 D. unifoliolata J. Léonard, 77 Didelta (Compositae), 137, 140 Didierea madagascarensis Bâillon (Didiereaceae), 241, 242 D. trollii Capuron & Rauh, 242 Digitalis purpurea L. (Scrophulariaceae), 147, 155, 157 Digitaria (Gramineae), 169, 202 D. adscendens (Kunth) Henrard, 254 D. ankaratrensis A. C a m u s , 239 D. argyrograpta (Nées) Stapf, 194 D.biformis Willd., 243 D. brazzae (Franchet) Stapf, 173 D. diagonalis (Nées) Stapf, 173,174, 194 D. humbertii A . C a m u s , 239 D. macroblephara (Hackel) Stapf, 126, 128 D. milanjiana (Rendle) Stapf, 101 D. monodactyla (Nées) Stapf, 194 D. pentzii Stent, 193 D. tricholaenoides Stapf, 194 D. uniglumis (Hochst. ex A . Rich.) Stapf: see D. diagonalis Diheteropogon (Gramineae), 129 D. amplectens (Nées) W . D . Clayton, 202
335
D. emarginatus (De Wild.) Robyns: see D. grandiflorus D. grandiflorus (Hackel) Stapf, 173 Dillenia ferruginea (Bâillon) Gilg (Dilleniaceae), 257 Dilobeia (Proteaceae), 232, 235, 236 Dimorphotheca (Compositae), 141 Diosma (Rutaceae), 132 Diospyros (Ebenaceae), 25, 74, 187, 235, 236, 241 D. abyssinica (Hiem) F. White, 109, 122, 129, 164, 167, 178, 187 D. aco*cksii (de Winter) de Winter, 141 D. austro-africana de Winter, 194, 195 ss. rugosa (E. Meyer ex A . D C . ) de Winter, 135 D. batocana Hiern, 93, 97,173 D. chevalieri D e Wild., 77 D. comorensis Hiern, 255 D. consolatae Chiov., 188,189 D. cornii Cbiov., 188,189 D. dichrophylla (Gandoger) de Winter, 135, 200, 201 D. diversifolia Hiern, 258 D. elliotii (Hiern) F. White, 176 D. feliciana Letouzey & F. White, 176 D. férrea (Willd.) Bakh., 77,109 D. gabunensis Giirke, 77, 181 D. galpinii Hiern, 201 D. glabra (L.) de Winter, 134 D. grex F. White, 172 D. heterotricha (B. L. Burtt) F. White, 172 D. hoyleana F . White, 77 D. inhacaenis F. White, 199 D. latispathulata H . Perrier, 242 D. longiflora Letouzey & F. White, 83 D. lycioides Desf., 98,137, 141,191,193,194,195, 202 D. melanida Poir., 258 D. mespiliformis Hochst. ex A . D C , 55, 81, 83, 90, 91, 95, 96, 99,105, 107, 108,117, 178,186,187, 206, 211 D. monbuttensis Gürke, 81 D. natalensis (Harv.) Brenan, 200 D. perrieri Jumelle, 241 D. pseudomespilus Mildbr., 74 D. quiloensis (Hiern) F . White, 90 D. ramulosa (E. Meyer ex A . D C . ) de Winter, 140 D . scabrida (Harv. ex Hiera) de Winter, 201 D. seychellarum (Hiern) Kostermans, 257 D. simii (Kuntze) de Winter, 201 D. squarrosa Klotzsch, 189 D. tesselaria Poir., 257 D. villosa (L.) de Winter, 96 D. wagemansii F. White, 172 D. whyteana (Hiern) F . White, 195,196, 201 Dipcadi (Liliaceae), 158 D. serotinum (L.) Medik., 152 D. thollonianum Hua, 100 Diplachne fusca (L.) P. Beauv. ex Stapf (Gramineae), 262 D. paucinervis (Nées) Stapf: see Odyssea paucinervis Diplorhynchus (Apocynaceae), 87 D. condylocarpon (Muell. Arg.) Pichón, 64, 96, 97, 99,173 Diplotaxis (Cruciferae), 158 D. tenuisiliqua Del., 227 Dirachma socotrana Schweinf. (Dirachmaceae), 111 Dirichletia glaucescens Hiern: see Carphalea glaucescens Discoclaoxylon occidentale (Muell. Arg.) Pax & Hoffm. (Euphorbiaceae), 253 Discoglypremna (Euphorbiaceae), 74 D. caloneura (Pax) Prain, 81, 253 Dissous canescens (Graham) Hook.f. (Melastomataceae), 266
336
Index ofplant names
Dissotis (Melastomataceae)—contd D. incana (E. Meyer ex Hochst.) Triana: see D. canescens D. rotundifolia (Smith) Triana, 265 Distemonanthus (Leguminosae: Caesalpinioideae), 74 Bobera glabra (Forssk.) Poir. (Salvadoraceae), 113, 114, 117, 188, 189 D. loranthifolia (Warb.) Harms, 114 Dodonaea madagascariensis Radlk. (Sapindaceae), 237, 238 D. viscosa Jacq., 109, 115, 137, 140, 251, 257 Dolichos lablab L . (Leguminosae: Papilionoideae), 252 Dombeya (Sterculiaceae), 236, 237 D. burgessiae Gerrard, 115 D. cymosa Harv., 200 D. goelzenii K . Schum., 122, 167 D . kirkii Masters, 182 D. mukole Sprague: see D. kirkii D. quinqueseta (Del.) Exell, 210 D . rotundifolia (Hochst.) Planchón, 95, 96, 193, 196, 201 D . shupangae K . Schum., 174 Dorstenia foetida Schweinf. (Moraceae), 113 D. gigas Schweinf. ex Balf.f., 255 D . gypsophila Lavranos, 116 Dovea mucronata Masters: see Chondropetalum mucronatum Dovyalis abyssinica (A. Rich.) Warb. (Flacourtiaceae), 121 Dracaena (Agavaceae), 235, 241 D. arbórea (Willd.) Link, 82 D. camerooniana Baker, 91 D. cinnabari Balf.f., 115, 246, 255 D. draco (L.) L., 246, 247, 251 D . ellenbeckiana Engl., 115 D. hookerana K . Koch, 200 D. ombet Kotschy & Peyr., 224, 246 D. refexa L a m . , 238 D. schizantha Baker, 115 Drakebrockmannia (Gramineae), 111 D . somalensis Stapf, 120 Drosanthemum luederitzii (Engl.) Schwantes (Aizoaceae), 142 D. paxianum (Schltr. & Diels) Schwantes: see D. luederitzii Drosera (Droseraceae), 84, 108 Dryopteris filix-mas (L.) Schott (Aspidiaceae), 156 D. parasitica (L.) Kuntze, 252 Drypetes floribunda (Muell. Arg.) Hutch. (Euphorbiaceae), 178 D. gerrardii Hutch., 129, 164, 167, 199 D. glabra (Pax) Hutch., 253 D . leonensis Pax, 82 D. parvifolia (Muell. Arg.) Pax & K . Hoffm., 178 D. principum (Muell. Arg.) Hutch., 253 Duboscia (Tiliaceae), 74 Dumoria africana (Pierre) Dubard: see Tieghemella africana D . heckelii A . Chev.: see T. heckelii Duosperma eremophilum (Milne-Redh.) Napper (Acanthaceae), 119,120 Duvalia (Asclepiadaceae), 113 Dypsis (Palmaceae), 232, 235 Ebenuspinnata L. (Leguminosae: Papilionoideae), 154 Eberlanzia spinosa Schwantes (Aizoaceae), 195 Ecbolium amplexicaule S. Moore (Acanthaceae), 114 E. revolutum C . B. Clarke, 114 Echidnopsis (Asclepiadaceae), 114, 116 Echinocarpus (Elaeocarpaceae), 235 Echinocloa colora (L.) Link (Gramineae), 108,191 E.pyramidalis (Lam.) Hitchco*ck & Chase, 100,108,265, 266 E. scabra (Lam.) R o e m & Schultes, 100, 265 E. stagnina auct. non (Retz.) P. Beauv.: see E. scabra
Echium (Boraginaceae), 158, 246, 251 E. boissieri Steud., 158 E. hypertropicum W e b b , 251 E. pomponium Boiss.: see E. boissieri E. stenosiphon W e b b , 252 E. vukanorum A . Chev., 252 Ectadium virgatum E . Meyer (Asclepiadaceae), 142 Ectropothecium (Hypnaceae), 81 Edithcolea (Asclepiadaceae), 114 Ehretia rígida (Thunb.) Druce (Boraginaceae), 140, 193, 194, 195, 201 E. teitensis Giirke, 114 Ehrharta (Gramineae), 134 E. erecta L a m . , 115 Eichhornia crassipes (Martius) Solms-Laub. (Pontederiaceae), 265 E. natans (P. Beauv.) Solms-Laub., 265 Ekebergia capensis Sparrman (Meliaceae), 74, 105,129, 200, 201 E. pterophylla ( C . D C . ) Hofmeyr, 96,196 E. senegalensis Adr. Juss.: see E. capensis Elaeis guineensis Jacq. (Palmaceae), 80, 83, 174,188 Elaeodendron buchananii (Loes.) Loes.: see Cassine buchananii E. orientale Jacq., 257, 258 Elaeophorbia (Euphorbiaceae), 54 E. drupifera (Thonn.) Stapf, 178 Eleocharis (Cyperaceae), 262 E. acutangula (Roxb.) Schultes, 265 Eleusine jaegeri Pilger (Gramineae), 130 Elionurus argenteus Nées (Gramineae), 101, 169,173,194, 196 E. hirtifolius Hackel, 210 E. royleanus Nées ex A . Rich., 252 E. tristis Hackel, 239 Elvira biflora (L.) D C . (Compositae), 252 Elymandra androphila (Stapf) (Gramineae), 84,174 Elytropappus (Compositae), 132 E. rhinocerotis Less., 32, 132, 139 Enantia kummeriae Engl. & Diels (Annonaceae), 187 Encephalartos (Zamiaceae), 199 E. altensteinii Lehm., 200 E. ferox Bertol.f., 200 E. hildebrandtii A . Braun & Bouché, 188 E. villosus Lemaire, 200 Endodesmia (Guttiferae), 74 Englerodendron (Leguminosae: Caesalpinioideae), 186 E. usambarense Harms, 187 Enneapogon (Gramineae), 221 E. brachystachyus (Jaub. & Spach) Stapf: see E. desvauxii E. cenchroides C . E . Hubbard., 145, 254 E. desvauxii P. Beauv., 195, 221 E. scaber Lehm., 221 Entada abyssinica Steud. ex A . Rich (Leguminosae: M i m o soideae), 85,174 E. africana Guill. & Perr., 107 E. mannii (Oliver) Tisserant, 81 E. pursaetha D C , 81 E. spicata (E. Meyer) Druce, 200, 201 Entandrophragma angolense (Welw.) C . D C . (Meliaceae), 77, 172,181 E. candollei Harms, 77 E. caudatum (Sprague) Sprague, 90, 99, 201 E. cylindricum (Sprague) Sprague, 77, 181 E. delevoyi D e Wild., 90, 96 E. excelsum (Dawe & Sprague) Sprague, 85,164 E. palustre Staner, 83 E. utile (Dawe & Sprague) Sprague, 75, 77, 82, 181
Index of plant names
Enterospermwn (Rubiaceae), 237 Entolasia imbricata Stapf (Gramineae), 100 Ephedra alata D C . (Ephedraceae), 220 E. altissima Desf., 152, 227 E. fragilis Dcst., 153,154 E. tilhoana Maire, 222 Ephippiandra (Monimiaceae), 232, 236 Ephippiocarpa (Apoeynaceae), 199 Equisetum ramosissimum Desf. (Equisetaceae), 222, 252 Eragrostis (Gramineae), 116,144, 193 E. atherstonei Stapf, 194 E. biflora Hackel, 193 E. capensis (Thunb.) Triri., 194 E. chalcantha Trin.: see E. racemosa E. chloromelas Steud., 194 E. ciliaris (L.) R . Br., 101, 193 E. curvula (Schrader) Nees, 195 E. cyperoides (Thunb.) P. Beauv., 142 E. decumbens Renvoize, 259 E. gummiflua Nees, 194 E. haraensis Chiov., 115 E. kohorica Quézel, 222 E. lateritica Bosser, 239, 243 E. lehmannianatiees, 141, 193, 194,195 E. micrantha Hackel, 194 E. nindensis Ficalho & Hiern, 142 E. obtusa M u n r o ex Ficalho & Hiern, 194,195 E. pollens Hackel, 193 E. papposa (Dufour) Steud., 222 E. plana Nees, 194 E. porosa Nees, 144 E. racemosa (Thunb.) Steud., 194,196 E. sclerantha Nees, 194 E. scotelliana Rendle, 84 E. superba Peyr., 194 E. tenuifolia (A. Rich.) Steud., 122,126 E. trémula (Lam.) Hochst. ex Steud., 209, 210, 211, 212, 213 Eremospatha (Palmaceae), 83 Erica (Ericaceae), 132,134,149,168,195 E. arbórea L . , 129, 147, 152, 157, 159, 216, 222, 246, 248, 249 E. caffraL., 134 E. caterviflora Salisb., 134 E. inconstans Zahlbr., 134 E. multiflora L., 154 E. scoparia L . ss. azorica (Hochst.) D . A . W e b b , 246, 247 E. umbellata L., 152 Erinacea anthyllis Link (Leguminosae: Papilionoideae), 156, 158 Eriobotrya japónica (Thunb.) Lindley (Rosaceae), 252 Eriobroma oblongum (Masters) Bodard: see Sterculia oblonga Eriocephalus (Compositae), 135, 137,139, 140, 193 E. racemosus L., 135 E. spinescens Burchell, 195 Eriochloa meyerana (Nees) Pilger (Gramineae), 267 Eriosema (Leguminosae: Papilionoideae), 243 Eriospermum abyssinicum Baker (Liliaceae), 100 Erismadelphus (Vochysiaceae), 74 Erodium glaucophyllum (Geraniaceae), 221 Eryngium ilicifolium L a m . (Umbelliferae), 229 E. tricuspidatum L., 152 Erysimum caboverdeanum (A. Chev.) Sund. (Cruciferae), 252
337
Erythrina abyssinica D C . (Leguminosae: Papilionoideae), 107, 174 E. baumii H a r m s , 173 E. caffra Thunb., 199 E. excelsa Baker, 181 E. sacleuxii H u a , 187 E. sigmoidea H u a , 211 Erythrochlamys (Labiatae), 111 E. spectabilis Giirke, 114 Erythrococca bongensis Pax (Euphorbiaceae), 182 E. menyharthii (Pax) Prain, 90 Erythrophleum africanum (Welw.) H a r m s (Leguminosae: Caesalpinioideae), 87,91,93,96, 97,105,106,173 E. guiñéense G . D o n : see E. suaveolens E. lasianthum Corbishley, 200 E. suaveolens (Guill. & Perr.) Brenan, 74, 91,178, 187, 188 Erythrophysa (Sapindaceae), 241 Erythrostictus (Liliaceae), 158 Erythroxylum acranthum Hemsley (Erythroxylaceae), 259 E. emarginatum Thonn., 99,109 E. lanceum Bojer, 255 E. platycladum Bojer, 243 Eucalyptus (Myrtaceae), 160,231 Euclea (Ebenaceae), 98,115,201 E. coriácea A . D C , 195 E. crispa (Thunb.) Gtirke, 137, 194,195 ss. ovata (Burchell) F . White, 193,195 E. divinorum Hiern, 115, 129,167 E. lancea Thunb., 135 E. natalensis A . D C , 99, 188,189, 200 ss. capensis F . White, 135 E. pseudebenus E . Meyer ex A . D C , 141,144 E. racemosa Murray, 135, 200 ss. schimperi ( A . D C . ) F . White, 115, 121, 129, 182, 188,224 E. schimperi ( A . D C . ) Dandy: see E. racemosa ss. schimperi E. tomentosa E . Meyer ex A . D C , 135 E. undulata Thunb., 135, 137, 140,141,193, 201 Eugenia (Myrtaceae), 236 E. capensis (Eckl. & Zeyh.) Sond., 202 E. leonensis Engl. & v. Brehm., 82 E. sp., 258 Eulalia villosa (Thunb.) Nees (Gramineae), 202 Euonymus latifolius (L.) Miller (Celastraceae), 151 Euphorbia (Euphorbiaceae), 50, 54, 111, 115, 116, 117, 137, 139, 147, 149, 174,198, 201, 228, 229, 238, 241, 242, 249 E. arbúsculo Balf.f., 255 E. avasmontana Dinter, 140,193 E. azorica Seub., 246 E. balsamifera Aiton, 207 E. beaumierana Hook.f. & Cosson, 225, 226, 227, 228 E. bellica Hiern, 144 E. bilocularis N . E . Br., 97 E. calycina N . E . Br.: see E. candelabrum E. candelabrum Trémaux ex Kotschy, 98, 99, 115, 128, 129, 182,188,207,210 E. clavarioides Boiss., 195 E. columnaris Bally, 116 E. conspicua N . E . Br., 90, 91 E. cunéala Vahl, 116 E. currorii N . E . Br., 140 E. ¿awei N . E . B r . , 182, 183 E. desmondii Keay & Milne-Redh., 62,109 E. dinteri Berger: see E. virosa E. echinus Hook.f. & Coss., 224, 225, 226,227, 228, 229
338
Index ofplant names
Euphorbia (Euphorbiaceae)—contd E. eduardoi Leach, 140 E. enterophora Drake, 241 E. evansii Pax, 201 E. grandicornis Goebel, 114, 188 E. grandidens H a w . , 137, 200, 201 E. grandis Lemaire, 115 E. gregaria Marloth, 140 E. guerichiana Pax, 140 E. gummifera Boiss., 140, 142 E. inaequilatera Sond., 194 E. ingens E . Meyer ex Boiss., 96, 201 E. kamerunica Pax, 109 E. mauritanica L., 135, 140, 195 E. mosaica Bally & S. Carter, 116 E. multiclava Bally & S. Carter, 116 E. nyikae Pax, 114,128,188 E. origanoides L . , 254 E. phillipsae N . E . Br., 113 E. poissonii Pax, 109 E. pyrifolia L a m . , 259 E. quinquecostata Volkens, 114 E. regis-jubae W e b b & Berth., 224, 225, 226, 228, 229 E. resinífera Berger, 225, 226, 227, 228 E. robecchii Pax, 114 E. scheffieri Pax, 114 E. schimperi Presl, 120 E. sepulta Bally & S. Carter, 116 E. socotrana Balf.f., 115 E. spiralis Balf.f., 255 E. stenoclada Bâillon, 242 E. striata Thunb., 194 E. subsalsa Hiern, 145 E. sudanica A . Chev., 109 E. tetrágono H a w . , 200, 201 E. tirucalliL., 128,201 E. triangularis Desf., 200, 201 E. tuckeyana Steud., 246, 251, 252 E. virosa Willd., 144 E. wakefieldii N . E . Br., 188 Euryops (Compositae), 135, 140 Eurypetalum (Leguminosae: Caesalpinioideae), 74 Eustachys paspaloides (Vahl) Lanza & Mattei (Gramineae), 126, 128,137, 194 Excoecaria bussei (Pax) Pax (Euphorbiaceae), 90 E. venenífera Pax, 188 Exotheca abyssinica Anderson (Gramineae), 169 fa*gara capensis Thunb.: see Zanthoxylum capense F. chalybea (Engl.) Engl.: see Z . chalybeum F. davyi I. Verdoorn: see Z davyi F. macrophylla (Oliver) Engl.: see Z . gilletii F. trijuga Dunkley: see Z . trijugum F. xanthoxyloides L a m . : see Z . xanthoxyloides fa*garopsis angolensis (Engl.) Dale (Rutaceae), 167 fa*gonia (Zygophyllaceae), 230 F. crética L . , 229 F. flamandii Battand., 222 F. glutinosa Del., 220 F. latifolia Del., 221 F. microphylla Pomel, 221 F. mollis DA., 22\ Farsetia (Cruciferae), 111 F. aegyptiaca Turra, 221 F. longisiliqua Decne., 115
F. stenoptera Hochst., 204 Faurea (Proteaceae), 237 F. forficuliflora Baker, 237, 238 F. saligna Harv., 93, 96, 105, 167 F. speciosa W e l w . , 99 Fedia (Valerianaceae), 158 Fegimanra (Anacardiaceae), 74 Felicia (Compositae), 141 F. filifolia (Vent.) Burtt Davy, 194, 196 F. muricata (Thunb.) Nees, 194 Feretia aeruginescens Stapf (Rubiaceae), 99 Fernandoa madagascariensis (Baker) A . Gentry (Bignoniaceae), 243 F. magnifica Seemann, 117, 187 Ferula (Umbelliferae), 228, 230 F. communis L . , 158,160 Festuca (Gramineae), 156, 169, 195 F. abyssinica Hochst. ex A . Rich., 211 F. camusiana St Yves, 239 F. caprina Nees, 196 F. costata Nees, 169 F. hystrix Boiss., 156 F triflora Desf., 157 Ficalhoa (Theaceae), 162 F. laurifolia Hiern, 164 Ficinia (Cyperaceae), 132 Ficus (Moraceae), 75, 98,129, 209, 210, 259 F. annobonensis Mildbr. & Hutch., 253 F. capensis Thunb., 91,105, 187, 196, 199, 251 F. carica L . , 252 F. congensis Engl., 91,181, 266 F. cordata Thunb., 140, 193 F. exaspérala Vahl, 83 F. fischeri W a r b . ex Mildbr. & Burret, 90, 98 F. glumosa Del., 105, 109 F. guerichiana Engl., 140, 193 F. ingens (Miq.) Miq., 96, 99, 117, 196, 222 F. lecardii W a r b . , 109 F. marmorata Bojer, 242 F. natalensis Hochst., 199, 200 F. populifolia Vahl, 207 F. pseudosycomorus Decne., 224 F. sagittifolia W a r b . ex Mildbr. & Burret, 75 F. salicifolia Vahl, 207, 222 F. socotrana Balf.f., 115, 255 F. soldanella'Waib., 196 F. sonderi Miq., 99, 196 F. sycomorus L . , 91, 95, 96, 105, 117, 141, 144, 201, 211, 222,251,252 F. teloukat Battand., 222 F. trichopoda Baker, 200 F. vallis-choudae Del., 187 F. verruculosa Warb., 266 F. vogelii (Miq.) Miq., 83 Filicium decipiens (Wight & A m . ) T h w . (Sapindaceae), 255 Fimbristylis pilosa Vahl (Cyperaceae), 178 Fingerhuthia africana L e h m . (Gramineae), 137, 195 Fissidens sciophyllus Mitten (Fissidentaceae), 182 Flacourtia flavescensWmà. (Flacourtiaceae): seeF. indica F. indica (Burm.f.) Merr., 178, 189, 241 Fleurydora felicis A . Chev. (Ochnaceae), 176 Foeniculum vulgare Miller (Umbelliferae), 158 Foetidia mauritiania L a m . (Foetidiaceae), 257 F. rodriguesiana Friedmann, 258 Foley ola (Cruciferae), 218
Index of plant names
Fomes annosus (Fries) Cooke (Polyporaceae), 78 Forgesia borbónica Pers. (Escalloniaceae), 258 Forsskâlea tenacissima L. (Urticaceae), 204, 221 Frángula alnus Miller: see Rhamnus frángula Frankenia (Frankeniaceae), 223, 266 F. corymbosa Desf., 229 F. laevis L., 230 F. portulacifolia Spreng., 254 Fraxinus angustifolia Vahl (Oleaceae), 149,150 F. xanthoxy bides Wáñ., 149,153, 156 Fredolia aretioides M o q . ex Coss. (Chenopodiaceae), 221 Freylinia oppositifolia Spin (Scrophulariaceae), 135 Friesodiebia obovata (Benth.) Verde: see Popowia obovata Fuirena pubescens (Poir.) Kunth (Cyperaceae), 100 F. umbellata Rottb., 84, 266 Funtumia africana (Benth.) Stapf (Apocynaceae), 74, 81, 187, 253 Furcraea foetida (L.) H a w . (Agavaceae), 257 F. gigantea Vent.: see F. foetida Gaertnera (Rubiaceae), 82 Gagea (Liliaceae), 158 Galenia (Aizoaceae), 140, 193 Galpinia (Lythraceae), 199 G. transvaalica N . E . Br., 200 Garcinia chromocarpa Engl. (Guttiferae), 83 G. echirensis Pellegr.: see G. chromocarpa G. livingstoneiT. Andersson, 91,117, 129, 201 G. polyantha Oliver: see G. smeathmannii G. punctata Oliver, 74, 83 G. smeathmannii (Planchón & Triana) Oliver, 82, 91 Gardenia (Rubiaceae), 242 G. imperialis K . Schum., 91 G. jovis-tonantis (Welw.) Hiern: see G. ternifolia G. lútea Fresen.: see G. ternifolia G. sokotensis Hutch., 107,109 G. ternifolia Schumach. & Thonn., 85, 107, 108, 174, 210, 211 Garuleum (Compositae), 140 Gasteria (Liliaceae), 137,140 Geigeria (Compositae), 194 G. alata ( D C . ) Benth. & Hook.f. ex Oliver & Hiern, 204 G. áspera Harv., 194 G. spinosa O . Hoffm., 144,145 Genista (Leguminosae: Papilionoideae), 147, 229 G. ferox Poir., 227 G. myriantha Bail, 153 G. saharae Coss. & Durieu, 220 G. retamoides Spach, 154 G. tricuspidata Desf., 157 Geopanax (Araliaceae), 257 Geophila (Rubiaceae), 75 Gerrardanthus lobatus (Cogn.) C . Jeffrey (Cucurbitaceae), 114 Geum syhaticum Pourret (Rosaceae), 157 Gilbertiodendron (Leguminosae: Caesalpinioideae), 74 G. bilineatum (Hutch. & Dalz.) J. Léonard, 77 G. brachystegioides (Harms) J. Léonard, 77 G. dewevrei (De Wild.) J. Léonard, 78, 79 G. ogoouense (Pellegr.) J. Léonard, 79 G. preussii (Harms) J. Léonard, 77 G. splendidum (A. Chev. ex Hutch. & Dalz.) J. Léonard, 77 Gilletiodendron glandulosum (Portères) J. Léonard (Leguminosae: Caesalpinioideae), 103 Givotia gosai Radcl.-Smith (Euphorbiaceae), 114 G. madagascariensis Bâillon, 241
339
Gladiolus (Iridaceae), 158 G. byzantinus Miller, 158 Globularia alypum L. (Globulariaceae), 153, 155, 222, 226, 229 G. amygdalifolia W e b b , 252 Glossonema boveanum (Decne.) Decne. (Asclepiadaceae), 204 Glumea ivorensis Aubrév. & Pellegr. (Sapotaceae), 76, 77 Gnidia (Thymelaeaceae), 194 G. glauca (Fresen.) G ü g , 130 G. kraussiana Meissner, 173,194 G. polycephala (C. A . Meyer) Gilg, 195 G. subcordata Meissner, 115 Gossweilerodendron (Leguminosae: Caesalpinioideae), 74 Gossypium hirsutum L . (Malvaceae), 252 G. somátense (Gürke) J. B . Hutch., 204 Grandidiera (Flacourtiaceae), 186 G. boivinii Jaub., 188 Grangeria borbónica L a m . (Chrysobalanaceae), 258 Greenwayodendron suaveolens (Engl. & Diels) Verde. (Annonaceae), 74 ss. usambaricum Verde, 187 Grewia (Tiliaceae), 98,113, 236, 242 G. avellana Hiern, 90 G. bicolor Juss., 182 G. burttii Exell, 97 G. carpinifolia Juss., 91,178 G. fallax K . Schum., 114, 128 G. flava DC, 96,140, 193 G. flavescens Juss., 90, 210, 211, 213 G. megalocarpa Juss., 176 G. mollis Juss., 174, 182,211 G. occidentalis L., 194, 195, 201 G. plagiophylla K . Schum., 189 G. robusta Burch., 201 G. similis K . Schum., 115,121,129,182 G. tembensis Fresen., 114,115 G. tenax (Forssk.) Fiori, 114, 213, 222 G. trichocarpa A . Rich., 129 G. truncata Masters, 189 G. villosa Willd., 95, 114, 176, 211, 251 Grielum (Neuradaceae), 137, 141 Griffonia simplicifolia (Vahl ex D C . ) Bâillon (Leguminosae: Caesalpinioideae), 178 Grimmia campestris Burchell ex Hooker (Grimmiaceae), 53 G. ovalis (Hedwig) Lindberg, 53 G. ovata Weber & M o h r : see G. ovalis Grossera (Euphorbiaceae), 74 Grubbia (Grubbiaceae), 132 Guarea cedrata (A. Chev.) Pellegr. (Meliaceae), 77, 82 G. thompsonii Sprague & Hutch., 77 Guibourtia copallifera Bennett (Leguminosae: Caesalpinioideae), 103, 109 G. demeusei (Harms) J. Léonard, 83 Guiera senegalensis J. F . G m e l . (Combretaceae), 203, 210, 212 Gymnorinorea: see Decorsella Gymnosiphon (Burmanniaceae), 75 Gyrocarpus americanus Jacq. (Hernandiaceae), 109, 242 Gyroptera (Chenopodiaceae), 111
Haematostaphis (Anacardiaceae), 103 H . barteri Hook.f., 105 Hagenia (Rosaceae), 162 H . abyssinica (Bruce) J. F . Gmelin, 47, 130, 161, 165, 166, 167 Hakea acicularis (Vent.) Knight (Proteaceae), 135
340
Index of plant names
Halimium (Cistaceae), 147 H . atlanticum Humbert & Maire, 157 H . halimiifolium (L.) Willk., 151 H . lasiocalycinum (Boiss. & Reuter) Maire, 152 H . libanotis Lange, 151 Halleria lucida L . , (Scrophulariaceae), 165,195,196, 201 Halocnemum strobilaceum (Pallas) M . Bieb. (Chenopodiaceae), 233,224,230 Haloxylon scoparium Pomel (Chenopodiaceae), 221, 228, 229, 230 Haplocarpha (Compositae), 194 H . scaposa Harv., 194 Haplocoelum foliolosum (Hiern) Bullock (Sapindaceae), 98, 99, 129,188 H . inoploeum Radlk., 188, 189 Harmsia (Sterculiaceae), 111 Harpachne (Gramineae), 111 Harpagophytum (Pedaliaceae), 241 Harpechloa faix (L.f.) Kuntze (Gramineae), 194 Harpephyllum (Anacardiaceae), 199 H . caffrum Bernh., 199, 200,201 Harrisonia abyssinica Oliver (Simaroubaceae), 189 Hartogia capensis L.f. (Celastraceae), 135 Harungana madagascariensis L a m . ex Poir. (Guttiferae), 80, 81, 83,105,173,236 Haworthia (Liliaceae), 137, 140 H . tesselata H a w . , 195 Haya Balf.f. (Caryophyllaceae), 111 Heberdenia bahamensis (Gaertner) Sprague (Myrsinaceae): see H . excelsa H . excelsa (Aitón) Banks ex D C , 246, 247 Heckeldora (Meliaceae), 74 Hederá helix L . (Araliaceae), 151, 156 Hedycaryopsis (Monimiaceae), 232, 236 Hedychium coronarium Koenig (Zingiberaceae), 236 Hedyotis adscensionis D C . (Rubiaceae), 254 H . arbórea Roxb., 254 Heeria argéntea (Thunb.) Meissner (Anacardiaceae), 134 H . concolor Presl ex Sond., 140 H . crassinervia (Engl.) Engl., 140,193 H . reticulata (Baker f.) Engl., 96, 97, 129, 189 Heisteria parvifolia Smith (Olacaceae), 253 Helianthemum (Cistaceae), 147 H . canariense Pers., 153, 227, 228 H. gorgoneum W e b b , 252 H . kahiricum Del., 221 H . lavandulifolium Miller, 154 H . pergamaceum Pomel, 230 Helichrysum (Compositae), 132,140, 168, 194, 237, 238, 239 H . dregeanum Sond. & Harv., 194
H. glumaceum D C , 116 H . latifolium (Thunb.) Less., 194 H . oreophilum Klatt, 194 H . rugulosum Less., 194 H . yuccifolium L a m . , 258 Heliophila (Cruciferae), 141 Heliotropium curassavicum L . (Boraginaceae), 145 H . rariflorum Stocks, 204 Heritiera littoralis Dryander (Sterculiaceae), 261, 262, 263, 264 H . utilis (Sprague) Sprague: see Tarrietia Milis Hermannia (Sterculiaceae), 135, 137, 140, 141, 143 H . betonicifolia Eckl. & Zeyh., 194 H . candidissima Spreng.f., 195 H . coccocarpa Kuntze, 194, 195 H. depressa N . E . Br., 194
Hernandia ovigera L . (Hernandiaceae), 257 H . voyroni Jumelle, 241 Heteromorpha (Umbelliferae), 237 H . arborescens (Sprengel) C h a m . & Schlechtd., 194 Heteropogon (Gramineae), 238 H . contortus (L.) P. Beauv. ex R o e m . & Schult., 193, 194, 195,196,202,208,239,243,252 Hexalobus monopetalus (A. Rich.) Engl. & Diels (Annonaceae), 109 Heywoodia lucens Sim (Euphorbiaceae), 199 Hibiscus (Malvaceae), 140, 220 H . asper Hook.f., 108 H . diversifolius Jacq., 265 H . marlothianus K . Schum., 195 H . micranthus L.f., 145 H . tiliaceus L., 188, 257, 264 Hildebrandtia (Convolvulaceae), 111 Hildegardia barteri (Masters) Kosterm. (Sterculiaceae), 79, 83 Hippobromus (Sapindaceae), 199 H . pauciflorus (L.f.) Radlk., 200 Hippocratea indica Willd. (Celastraceae), 99 H . parviflora N . E . Br., 90 Hippocrepis (Leguminosae: Papilionoideae), 228 Hirtella (Chrysobalanaceae), 186 Holarrhena floribunda (G. D o n ) Dur. & Schinz (Apocynaceae), 81,83 Holoptelea grandis (Hutch.) Mildbr. (Ulmaceae), 79, 81, 181 Homalium (Flacourtiaceae), 83 H. dentatum (Harv.) W a r b . , 199 Hornería (Iridaceae), 141 Hoodia (Asclepiadaceae), 139 H. currori (Hook.) Decne., 142, 145 Hornea mauritania Baker (Sapindaceae), 257 Huernia (Asclepiadaceae), 139 Humbertochloa bambusiuscula A. C a m u s & Stapf (Gramineae), 242 Hyaenanche globosa (Gaertn.) Lambert (Euphorbiaceae), 134 Hydrilla verticillata Caspary (Hydrocharitaceae), 265 Hydrocotyle (Umbelliferae), 236 Hydrodea bossiana Dinter: see Mesembryanthemum cryptanthum H . cryptantha (Hook.f.) N . E . Br.: see M . cryptanthum Hygrophila auriculata (Schumach.) Heine (Acanthaceae), 108 Hylodendron (Leguminosae: Caesalpinioideae), 74 Hymenaea (Leguminosae: Caesalpinioideae), 186 H . verrucosa Gaertn., 188 Hymenocardia acida Tul. (Euphorbiaceae), 85, 97, 99, 105, 173, 174 H . ulmoides Oliver, 83, 200 Hymenocoleus (Rubiaceae), 75 Hymenodictyon floribundum (Steud. & Hochst.) B . L . Robinson (Rubiaceae), 81, 82, 83 H . parvifolium Oliver, 114 Hymenostegia (Leguminosae: Caesalpinioideae), 74 H . afzelii (Oliver) Harms, 77, 80 H . laxiflora (Benth.) H a r m s , 172 Hyophorbe (Palmaceae), 257 H . verschaffeltii H . A . Wendl., 258 Hyoscyannus muticus L . (Solanaceae), 219 Hyparrhenia (Gramineae), 50, 97, 129, 169, 189, 194, 209, 210, 238 H . anthistirioides (Hochst.) Andersson ex Asch. & Schweinf., 108,210,212 H . bracteata (Willd.) Stapf, 100 H . confinis (A. Rich.) Stapf, 173, 209 H . cyanescens (Stapf) Stapf, 107
Index ofplant names
H. H. H. H. H. H.
cymbaria (L.) Stapf, 128, 243 dichroa (Steud.) Stapf, 101 diplandra (Hackel) Stapf, 85, 100, 173 familiaris (Steud.) Stapf, 85,173 filipéndula (Hochst.) Stapf, 128,174, 202, 210 hirta (L.) Stapf, 128, 135, 137, 193, 195, 196, 208, 211, 252, 255 H . lecomtei (Franchet) Stapf: see H. newtonii H . multiplex (Hochst. ex A . Rich) Andersson ex Stapf, 211 H . mutica W . D . Clayton, 84 H . newtonii (Hackel) Stapf, 100, 101, 174, 239 H . nyassae (Rendle) Stapf, 85, 239 H . pachystachya Stapf: see H. diplandra H . papillipes (Hochst.) Andersson ex Asch. & Schweinf., 208 H . petiolata Stapf, 212 H . pseudocymbaria (Steud.) Stapf: see H . anthistirioides H . rufa (Nées) Stapf, 85,108, 174, 209, 239, 243 H . ruprechtii Fourn.: see Hyperthelia dissoluta H . schimperi (Hochst. ex A . Rich.) Andersson, 243 H . subplumosa Stapf, 32, 85 Hypericum lalandii Choisy (Guttiferae), 266 H . lanceolatum L a m . : see H . revolutum H. revolutum Vahl, 166, 258 H . roeperanum Schimp. ex A . Rich., 74 Hyperthelia dissoluta (Nées ex Steud.) W . D . Clayton (Gramineae), 101, 107,128, 174, 239, 243 Hyphaene (Palmaceae), 216, 219, 220 H . benguellensis W e l w . : see H . ventricosa H . compressa H . A . W e n d l , 188, 189 H . coriácea Gaertner, 123 H . natalensis Kuntze, 201 H . petersiana Klotzsch: see H . ventricosa H . shatan Bojer, 243 H . thebaica (L.) Martius, 55,107,108, 219, 220, 223 H . ventricosa Kirk, 95 Hypodaphnis (Lauraceae), 74 Hypoestes verticillaris (L.f.) R . Br. (Acanthaceae), 90 Hypolytrum (Cyperaceae), 75 Hypoxis angustifolia L a m . (Hypoxidaceae), 100 H . rigidula Baker, 194 H. rooperi S. Moore, 194 Icomum lineare Burkill (Labiatae), 100 Ifloga spicata (Forssk.) Schultes Bip. (Compositae), 220 Ilex aquifolium L . (Aquifoliaceae), 149,151, 156,157 /. canariensis Poir., 246 /. mitis (L.) Radlk., 74, 91, 122, 135, 165, 167, 181, 195, 196,237, 238 /. perado Aiton, 246 ss. azorica (Loes.) Tutin, 247 ss. platyphylla (Webb & Berth.) Tutin, 246 Imbricaría seychellarum Oliver (Sapotaceae), 257 Impatiens (Balsaminaceae), 165,167,236,238 /. irvingii Hook.f. ex Oliver, 265 Imperata cylindrica (L.) P. Beauv. (Gramineae), 50, 83, 85, 174, 224,236,239,243 Indigofera (Leguminosae: Papilionoideae), 111, 140, 243 /. alternons D C , 194 /. cordifolia Heyne ex Roth, 204 /. cunenensis Torre, 142 /. daleoides Benth., 144 /. disjuncta J. B . Gillett, 204 /. rhynchocarpa W e l w . ex Baker, 98 /. rostrata Bolus, 194
341
/. senegalensis L a m . , 204 /. sokotrana Vierh., 115 /. spinosa Forssk., 116, 119 /. subcorymbosa Baker, 98 /. teixeirae Torre, 145 Indokingia (Araliaceae), 257 Inhambanella henriquesii (Engl. & W a r b . ) Dubard (Sapotaceae), 187, 200 Intsia bijuga (Colebr.) Kuntze (Leguminosae: Caesalpinioideae), 257 Iphiona (Compositae), 242 Ipomoea (Convolvulaceae), 111,114, 265 /. crassipes H o o k . , 194 /. pes-caprae (L.) R . Br., 253, 254, 257, 258 /. sultani Chiov., 116 /. verbascoidea Choisy, 211 Iris (Iridaceae), 158 Irvingia gabonensis (Aubry-Lecomte ex O'Rorke) Bâillon (Irvingiaceae), 253 /. smithii Hook.f., 83 Isalus (Gramineae), 238 Ischaemum (Gramineae), 189 Isoberlinia (Leguminosae: Caesalpinioideae), 54, 58, 59, 61, 92, 96,102,103,106,107,176 /. angolensis (Welw. ex Benth.) Hoyle & Brenan, 87, 91, 92, 105,106 /. doka Craib & Stapf, 105, 106, 108 /. scheffieri (Harms) Greenway, 187 /. tomentosa (Harms) Craib. & Stapf: see /. angolensis Isolona heinsenii Engl. & Diels (Annonaceae), 187 Jardinea congoensis (Hackel) Franchet (Gramineae), 84 J. gabonensis Steudel, 84 Jasminum fluminense Vrell. (Oleaceae), 182 J.fruticans L., 153,154,158 J. mauritianum Bojer ex D C : see/, fluminense Jatropha(Euphorbiaceae), 111,242 J. curcas L . , 252, 255 / . glandulosa Vahl: see / . pelargoniifolia J. gossypiifolia L . , 252 J. pelargoniifolia Court»., 115 J. unicostata Balf.f., 255 J. villosa (Forssk.) Muell. Arg.: see J. pelargoniifolia Jubaeopsis (Palmaceae), 199 Julbernardia (Leguminosae: Caesalpinioideae), 92, 96, 97,106 J. globiflora (Benth.) Troupin, 92, 93, 99 J. magnistipulata (Harms) Troupin, 187, 188 J. paniculata (Benth.) Troupin, 91, 92, 97 J. pellegriniana Troupin, 77 J. seretii (De Wild.) Troupin, 77, 78, 79, 85 Juncus acutus L . (Juncaceae), 160, 219,246,266 J. arabicus (Aschers. & Buchenau) A d a m s o n , 224 / . bufonius L . , 222 J. effusus L., 191 J. maritimus L a m . , 222, 223, 266, 267 Juniperus (Cupressaceae), 115 / . brevifolia (Seub.) Antoine, 246, 247 J. cedrus W e b b & Berth., 246 J. communis L . , 153,158 J. oxycedrus L., 149,151,152, 154, 156, 158, 226 J.phoenicea L., 146, 149, 150, 151, 152, 153, 154, 155, 157, 159,226,229,248,249 / . procera Hochst. ex Endl., 51, 115, 122, 130, 161, 164, 165,166, 167 J. thurifera L., 146,149,150, 152,153, 155, 156
342
Index ofplant naines
Justiciaflava(Forssk.) Vahl (Acanthaceae), 182 Kaempferia rosea Schweinf. ex Benth. & Hook.f. (Zingiberaceae), 90 Kalanchoe (Crassulaceae), 114, 115, 129, 201, 236, 238, 241, 242 K. robusta Balf.f., 255 Kanahia (Asclepiadaceae), 111 Kaokochloa (Gramineae), 137 K. nigrirostis de Winter, 142 Kaoue stapfiana (A. Chev.) Pellegr. (Leguminosae: Caesalpinioideae), 77 Kedrostis gijef(J. F. Gmelin) C . Jeffrey (Cucurbitaceae), 114 Kelleronia (Zygophyllaceae), 111 K. quadricomuta Chiov., 116 Khaya anthotheca (Welw.) C D C . (Meliaceae), 81, 172, 181 K. comorensis Legris n o m . nud., 255 K. grandifoliola C D C , 79, 82, 181 K. nyasica Stapf ex Baker f., 91, 117,186 K. senegalensis (Desr.) Adr. Juss, 105, 178, 210, 211, 214 Kigelia africana (Lam.) Benth. (Bignoniaceae), 91, 95, 105, 117, 209 Kigelianthe madagascariensis (Baker) Sprague: see Fernandoa madagascariensis Kiggelaria (Flacourtiaceae), 162 K. africana L., 135, 165, 195, 199 Kirkia acuminata Oliver (Simaroubaceae), 95, 96, 99 K. wilmsii Engl., 96 Kissenia (Loasaceae), 113 K. capensis Endl., 113 Klainedoxa gabonensis Pierre ex Engl. (Irvingiaceae), 80, 172, 181 Kleinia (Compositae), 116, 139 K. cliffordiana (Hutch.) C . D . A d a m s , 109 K. kleinioides (Schultz-Bip.) M . R . F. Taylor, 120 K. scottii (Balf.f.) Chiov., 255 Koeleria (Gramineae), 169 K. pubescens (Lam.) P . Beauv., 230 K. vallesiana (Honck.) Bertol., 230 Kohautia amatymbica Eckl. & Zeyh. (Rubiaceae), 194 K. áspera (Roth) Bremek., 113 Kotschya africana Endl. (Leguminosae: Papilionoideae), 266 Kyllinga (Cyperaceae), 126 K. alba Nees, 116 K. erecta Schumach., 100 Lablab niger Medic: see Dolichos lablab L. purpureus (L.) Sweet: see D . lablab Laburnum platycarpum Maire (Leguminosae: Papilionoideae), 227 Lachanodes (Compositae), 254 L. arbórea (Roxb.) R . B. Nordenstam, 254 Lachenalia (Liliaceae), 141 Lachnocapsa (Cruciferae), 111 Lagarosiphon (Hydrocharitaceae), 265 Lagenantha nogalensis Chiov. (Chenopodiaceae), 120 Laguncularia racemosa Gaertner (Combretaceae), 261, 262 Lamarckia áurea (L.) Moench (Gramineae), 227 Landolphia (Apocynaceae), 241 L. camptoloba (K. Schum.) Pichón, 173 L. parvifolia K . Schum., 99 Lannea alata (Engl.) Engl. (Anacardiaceae), 114 L. amaniensis Engl. & K . Krause: see L. welwitschii L. antiscorbutica (Hiern) Engl., 173 L. discolor (Sond.) Engl., 96, 97, 98, 99, 196 L. fruticosa (Hochst. ex A . Rich.) Engl., 209
L. humilis (Oliver) Engl., 105, 116, 208, 209, 212 L. microcarpa Engl. & K . Krause, 107 L . schimperi (Hochst. ex A . Rich.) Engl., 105, 107 L . stuhlmannii (Engl.) Engl., 95, 129,189, 201 L . triphylla (Hochst. ex A . Rich.) Engl., 114 L. welwitschii (Hiern) Engl., 187, 253 Lantana (Verbenaceae), 189 L. cámara L., 236, 252 Lapeirousia (Iridaceae), 141 Lasiochloa (Gramineae), 134 L . echinata (Thunb.) Adamson, 135 Lasiocorys argyrophylla Vatke (Labiatae), 116 Lasiurus hirsutus (Forssk.) Boiss. (Gramineae), 221, 224 Latania commersonii J.F. Gmelin (Palmaceae): see L. lontaroides L . lontaroides (Gaertner) H . E . Moore, 257 L. verschaffeltii Lemaire, 258 Lathyrus (Leguminosae: Papilionoideae), 157 Launaea arborescens (Battand.) Maire (Compositae), 224, 229 L . chevalieri O . Hoffm. & Muschler, 204 Laurophyllus capensis Thunb. (Anacardiaceae), 134 Laurus azorica (Seub.) Franco (Lauraceae), 246, 247,249 L. nobilis L., 147, 149, 150, 226 Lavandula (Labiatae), 147, 221, 228, 229 L . coronopifolia Poir.: see L. stricta L. dentata L., 152,154, 227, 252 L. maroccana Murbeck, 228 LmultifidaL., 153, 154,227 L . pubescens Decne., 211, 222 L . rotundifolia Benth., 252 L. stoechasL., 151, 152 L. stricta Del., 219 Lebeckia (Leguminosae: Papilionoideae), 140 L . macrantha Harv., 193 Lebrunia bushaie Staner (Guttiferae), 79, 85 Lecaniodiscus fraxinifolius Baker (Sapindaceae), 91,117, 129 Leersia hexandra Swartz (Gramineae), 100, 266 Lemna (Lemnaceae), 265 L. perpusilla Torrey, 265 Leonotis mollissima Gürke (Labiatae), 122 Lepidopilum callochlorum C . Mueller ex Broth. (Daltoniaceae), 81 Lepidotrichilia volkensii (Gürke) Leroy (Meliaceae), 122, 167 Lepisanthes senegalensis (Poir.) Leenhouts: see Aphonia senegalensis Leptadenia pyrotechnica (Forssk.) Decne. (Asclepiadaceae), 204, 206, 207, 212, 213, 219, 220 L . reticulata Wight, 243 Leptaspis (Gramineae), 75 Leptochloa uniflora Hochst. ex A . Rich. (Gramineae), 90 Leptolaena bojerana (Bâillon) Cavaco (Sarcolaenaceae), 237 L . pauciflora Baker, 237 Leptothrium senegalense (Kunth) W . D . Clayton (Gramineae), 119,121 Lepturus (Gramineae), 266 Leucadendron (Proteaceae), 132,134 L . argenteum (L.) R . Br., 134 L . concinnum R . Br.: seeL. procerum L. eucalyptifolium E . M e y . ex Meissner, 134 L. nobileJ. M . Williams, 134 L . procerum (Salisb. ex Knight) J . M . Williams, 134 L . sabulosum Salter, 134 L. salicifolium J. M . Williams, 134,135 L. salignum R . Br., 135 Leucaena glauca auct. (Leguminosae: Mimosoideae): see L. leucocephala
Index ofplant names
L. leucocephala (Lam.) de Wit, 257 Leucojum (Amaryllidaceae), 158 Leucosidea (Rosaceae), 162 L. sericea Eckl. & Zeyh., 195, 196 Leucospermum (Proteaceae), 132, 134 L. conocarpodendron (L.) Buek, 134 Leucosphaera (Amaranthaceae), 137 L. bainesii (Hook.f.) Gilg, 191, 193, 267 Leuzea conifera (L.) D C . (Compositae), 155, 226 Librevillea (Leguminosae: Caesalpinioideae), 74 Lightfootia (Campanulaceae), 135, 140 Ligustrum robustum Blume (Oleaceae), 257 L. vulgareL., 151 Limnophyton obtttsifolium (L.) Miq. (Alismataceae), 265 Limoniastrum feei (de Gir.) Battand. (Plumbaginaceae), 221 L. guyonianum Durieu, 223 L. ifniense (Caball.) Font Quer, 223 L. monopetalum (L.) Boiss., 230 Limonium (Plumbaginaceae), 246 L cymuliferum (Boiss.) Sauvage & Vindt, 230 L. fallax (Wangerin) Maire, 229 L. pruinosum (L.) Kuntze, 223 Linaria (Scrophulariaceae), 147 L. sagittata Steud., 229 Lindackeria dentata (Oliver) Gilg (Flacourtiaceae), 81 Linociera foveolata (E. Meyer) Knobl.: see Chionanthus foveolatus Linum villarianum Pau (Linaceae), 157 Lippia ukambensis Vatke (Verbenaceae), 122 Lithops (Aizoaceae), 140, 142 Lobelia (Campanulaceae), 53 L. bambuseti R . E . Fries, 167 L. barnsii Exell, 253 Lobostemon (Boraginaceae), 132 Lochia (Caryophyllaceae), 111 Lodoicea (Palmaceae), 257 L. maldivica (J. F . Gmelin) Pers., 257 Loesenera (Leguminosae: Caesalpinioideae), 74 Loewia (Turneraceae), 111 Lonchocarpus bussei H a r m s (Leguminosae: Papilionoideae), 189 L. capassa Rolfe, 95, 96, 201 L. laxiflorus Guill. & Perr., 105,107, 211 L. nelsii (Schinz) Schinz ex Heering & G r i m m e , 90 Lonicera arbórea Boiss. (Caprifoliaceae), 156 L. etrusca G . Santi, 151,157 L. pyrenaica L., 158 Lophiocarpus polystachyus Turcz. (Chenopodiaceae), 145 Lophira ¿lata Banks ex Gaertner f. (Ochnaceae), 75, 77, 78 L. lanceolata V a n Tiegh. ex Keay, 85,105 Loranthus (Loranthaceae), 95 Lotononis tenuis Baker (Leguminosae: Papilionoideae), 145 Lotus (Leguminosae: Papilionoideae), 246 L. arabicus L., 144, 204 L. glinoides Delarbre, 252 L. mossamedensis W e l w . ex Baker: see L. arabicus Loudetia (Gramineae), 129, 238 L. arundinacea (Hochst. ex A . Rich.) Steud., 85, 173,174 L. demeusii (De Wild.) C . E . Hubbard, 173 L. filifolia Schweick. ss. humbertiana A. C a m u s , 243 L. kagerensis (K. Schum.) C . E . Hubbard ex Hutch., 84 L. phragmitoides (Peter) C . E . Hubbard, 85, 265, 266 L. simplex (Nées) C . E . Hubbard, 40, 50, 52, 84, 85, 100, 101,169,173, 202, 209, 210
343
ss. stipoides (Hackel) Bosser, 239, 243 L. togoensis (Pilger) C . E . Hubbard, 208, 209, 210 Loudetiopsis ambiens (K. Schum.) Conert (Gramineae), 84 L. glabrata (K. Schum.) Conert, 84 Lovoa swynnertonii Baker f. (Meliaceae), 186,187 L. trichilioides Harms, 77, 172 Loxostylis (Anacardiaceae), 199 Ludia (Flacourtiaceae), 186 L. mauritiana Gmelin, 189 L. sessiliflora L a m . : see L. mauritiana Ludwigia (Onagraceae), 265 L. erecta (L.) Hara, 265 L. leptocarpa (Nutt.) Hara, 265 L. octovalvis (Jacq.) Raven, 265 L. stolonifera (Guill. & Perr.) Raven, 265 Lumnitzera racemosa Willd. (Combretaceae), 261, 263, 264 Lupinuspilosus L . (Leguminosae: Papilionoideae): seeL. varius L. varius L., 147 Luzula fosteri (Smith) D C . (Juncaceae), 155 L. multiflora (Retz.) Lej., 156 L. sylvatica (Hudson) Gaudin, 156 Lycium (Solanaceae), 140, 143, 223 L. austrinum Miers, 201 L. decumbens Welw. ex Hiern, 145 L. europaeum L., 116 L. intricatum Boiss., 152, 223, 224, 228, 229, 266 L. tetrandrum L.f., 144 Lycopodium affine Bory (Lycopodiaceae), 84, 108 L. carolinianum L., 84 L. cernuum L., 84 L. mildbraedii Hert., 82 Lygeum (Gramineae), 230 L. spartum L., 149, 226, 229, 230 Lytanthus amygdalifolius (Webb) Wettst.: see Globularia amygdalifolia Macaranga (Euphorbiaceae), 236 M . capensis (Bâillon) T . R . Sim, 187 M . kilimandscharicaYax, 181 M . monandra Muell. Arg., 81, 181 M . pynaertii D e Wild., 181 M . schweinfurthii Pax, 181 M . spinosa Muell. Arg., 81 Maerua (Capparidaceae), 111,219 M . angolensis D C , 87,105,145 M . crassifolia Forssk., 119, 204, 206, 207, 212, 219 M . denhardtiorum Gilg, 114 M . filiformis Drake, 242 M . mildbraedii Gilg & C . Benedict: see M. triphylla M . subcordata (Gilg) D e Wolf, 114 M . triphylla A . Rich., 182 Maesa lanceolata Forssk. (Myrsinaceae), 211, 253 Maesopsis eminii Engl. (Rhamnaceae), 81, 181 Magnistipula butayei D e Wild. (Chrysobalanaceae), 74 ss. greenwayi (Brenan) F . White, 187 Malacantha alnifolia (Baker) Pierre (Sapotaceae), 83, 187 Malcolmia aegyptiaca Sprengel (Cruciferae), 220 Malus domestica Borkh. (Rosaceae), 252 M . sylvestris Mill.: see M . domestica Mammea (Guttiferae), 235 M . africana Sabine, 253 Mangifera (Anacardiaceae), 189 M . indica L . , 174, 252 Manilkara (Sapotaceae), 187 M . concolor (Harv. ex C . H . Wright) Gerstner, 96,199
344
Index ofplant names
Manilkara (Sapotaceae)—contd M . discolor (Sond.) J. H . Hemsley, 200 M . mochisia (Baker) Dubard, 91, 188, 189 M . obovata (Sabine & G . D o n ) J. H . Hemsley, 81, 83, 167, 178 M . sansibarensis (Engl.) Dubard, 187, 188,189 M . sulcata (Engl.) Dubard, 116, 188 Mansonia altissima (A. Chev.) A . Chev. (Sterculiaceae), 79 Mapania (Cyperaceae), 75, 77 Maprounea africana Muell. Arg. (Euphorbiaceae), 97, 105, 173, 174 Maranthes glabra (Oliver) Prance (Chrysobalanaceae), 77, 85 M . goetzeniana (Engl.) Prance, 186,187 M . polyandra (Benth.) Prance, 83, 85, 105 Marantochloa (Marantaceae), 75 Marattia (Marattiaceae), 82 Margaritaria discoidea (Bâillon) Webster: see Phyllanthus discoideus Mariscus deciduus C . B. Clarke (Cyperaceae), 100 Markhamia acuminata (Klotzsch) K . Schum. (Bignoniaceae), 90 M . hildebrandtii (Baker) Sprague, 167 M . obtusifolia (Baker) Sprague, 90,95 Marquesia (Dipterocarpaceae), 173 M. acuminata (Gilg) R . E . Fries, 90, 173 M . macroura Gilg, 90, 91, 93, 173 Mascarena verschaffeltii (H. A . Wendl.) L . H . Bailey: see Hyophorbe verschaffeltii Mathurina penduliflora Balf.f. (Turneraceae), 258 Matthiola kralikii Pomel (Cruciferae), 229 May tenus (Celastraceae), 201 M . acuminata (L.f.) Loes., 135, 195 M . heterophylla (Eckl. & Zeyh.) N . Robson, 115, 135, 193, 195 M . linearis (L.f.) Marais, 201, 243 M . oleoides (Lam.) Loes., 134, 135 M . polyacantha (Sond.) Marais, 195 M . senegalensis (Lam.) Exell, 85, 91, 174, 189, 207, 222, 228, 259 M . undata (Thunb.) Blakelock, 195 Medemia argun (Martius) Württemb. ex H . A . Wendl. (Palmaceae), 218 M . nobilis Gallerand, 243 Medinilla (Melastomataceae), 236 Medusagyne oppositifolia Baker (Medusagynaceae), 257 Megalochlamys (Acanthaceae), 113 Megaloprotachne albescens C . E . Hubbard (Gramineae), 193 Megistostegium (Malvaceae), 232, 242 Melanodendron (Compositae), 254 M . integrifolium (Roxb.) D C , 254 Melanthera scandens (Schumach. & Thonn.) Roberty (Compositae), 265 Melastomastrum segregatum (Benth.) A . & R . Fernandes (Melastomataceae), 265 Melhania melanoxylon Aiton: see Trochetia melanoxylon Melia volkensii Giirke (Meliaceae), 114 Melinis minutiflora P. Beauv. (Gramineae), 252, 254 Mellissia (Solanaceae), 254 M . begoniifolia (Roxb.) Hook.f., 254 Memecylon (Melastomataceae), 187 M . eleagni Blume, 257 M . sansibaricum Taubert, 188 M . sapinii D e Wild., 173 Merremia multisecta Hallier f. (Convolvulaceae), 142 Merxmuellera (Gramineae), 134 M . disticha (Nées) Conert, 139
M . macowanii (Stapf) Conert, 239 M . stricta (Schrader) Conert, 139 Mesanthemum radicans (Benth.) Koern. (Eriocaulaceae), 84 Mesembryanthemum (Aizoaceae), 266 M . cryptanthum Hook.f., 142, 254 Mesogyne henriquesii Engl. (Moraceae), 253 Metalasia (Compositae), 132,134 M . muricata (L.) Less., 134,135 Metrosideros angustí folia (L.) Smith (Myrtaceae), 135 Michelsonia microphylla (Troupin) H a u m a n (Leguminosae: Caesalpinioideae), 78,79 Microberlinia bisulcata A . Chev. (Leguminosae: Caesalpinioideae), 77 Microchloa coffra Nées (Gramineae), 194 M . indica (L.f.) P. Beauv., 116, 213 M . kunthii Desv., 126,128, 209 Micromeria (Labiatae), 249 M . forbesii Benth., 249 Mikania cordata (Burm.f.) B. L . Robinson (Compositae), 265 Mildbraediodendron excelsum Harms (Leguminosae: Caesalpinioideae), 181 Milium vernale M . Bieb. (Gramineae), 157 Millettia grandis (E. M e y . ) Skeels (Leguminosae: Papilionoideae), 199,200 M . sutherlandii Harv., 199 M . thonningii (Schumach. & Thonn.) Baker, 178 M . usaramensis Taubert, 189 Mimetes (Proteaceae), 134 M . fimbrifolius Salisb. ex Knight, 134 Mimosa pigra L . (Leguminosae: Mimosoideae), 266 Mimusops aedificatoria Mildbr. (Sapotaceae), 187 M . caffra E . M e y . ex A . D C , 200 M . maxima (Lam.) Vaughan, 258 M . obovata Sond., 200 M . petiolaris ( D C . ) Dubard, 258 M . zeyheri Sond., 91, 99, 196 Miscanthus (Gramineae), 55 M . teretifolius (Stapf) Stapf, 100 M . violaceus (K. Schum.) Pilger, 265, 266 Mitolepis (Asclepiadaceae), 111 M . intricata Balf.f., 115 Mitragyna ciliata Aubrév. & Pellegr. (Rubiaceae), 83 M . inermis (Willd.) Kuntze, 105, 107, 108 M . rubrostipulata (K. Schum.) Havil., 85,164 M . stipulosa (DC.) Kuntze, 83, 91, 181 Molinaea sp. (Sapindaceae), 258 Monadenium invenustum N . E . Br. (Euphorbiaceae), 114 Monanthes (Crassulaceae), 246 Monanthotaxis fornicata (Bâillon) Verde. (Annonaceae), 189 Monechma (Acanthaceae), 140 M . genistifolium C. B . Clarke, 267 M . tonsum P. G . Meyer, 267 Monelytrum (Gramineae), 137 Monocyclanthus (Annonaceae), 74 Monocymbium ceresiiforme (Nées) Stapf (Gramineae), 84, 85, 100, 101, 169,173, 194, 196 Monodiella (Gentianaceae), 218 Monodora myristica (Gaertner) Dunal (Annonaceae), 181, 253 Monopetalanthus (Leguminosae: Caesalpinioideae), 74 M . compactus Hutch. & Dalz., 77 M . hedinii (A. Chev.) Pellegr., 77 M . richardsiae J. Léonard, 91 M . trapnellii J. Léonard, 91 Monotes (Dipterocarpaceae), 87, 99, 106, 176 M . caloneurus Gilg, 174
Index of plant names
M . dasyanthus Gilg, 173 M . kerstingii Gilg, 105 M. mutetetwa Duvign., 174 Monotheca buxifolia (Falcolner) A . D C : see Sideroxylon buxifolium Monsonia ignorata M e r x m . & A . Schreiber (Geraniaceae), 142 M. nivea W e b b , 220 M. senegalensis Guill. & Perr., 145 Montinia caryophyllacea Thunb. (Montiniaceae), 137, 193, 201 Moraea natalensis Baker (Iridaceae), 100 Moricandia arvensis (L.) D C . (Cruciferae), 221 Morinda asteroscepa K . Schum. (Rubiaceae), 187 Moringa (Moringaceae), 111, 242 M. ovalifolia Dinter & A . Berger, 140 M. peregrina (Forssk.) Fiori, 224 Monis láctea (Sim) Mildbr. (Moraceae): see M. mesozygia M. mesozygia Stapf ex A . Chev., 79, 81, 82, 105, 178, 181, 199 Mucuna sloanei Fawcett & Rendle (Leguminosae: Papilionoideae), 253 Mundulea phylloxylon R . Viguier (Leguminosae: Papilionoideae), 238 M. sericea (Willd.) A . Chev., 210 Muraltia (Polygalaceae), 132, 134 Musanga cecropiodes R . Br. (Moraceae), 25, 80, 81, 83,181, 253 M. leo-errerae H a u m a n & J. Léonard, 85 Muscari (Liliaceae), 158 Myrianthus arboreus P . Beauv. (Moraceae), 81 M . holstii Engl, 164,187 Myrica (Myricaceae), 237 M. faya Aiton, 246, 247, 248 Myrmecosicyos (Cucurbitaceae), 111 Myrothamnus (Myrothamnaceae), 238 M. flabellifolius (Sond.) W e l w . , 23, 99, 238 M. moschatus Bâillon, 238 Myrsine africana L . (Myrsinaceae), 135, 195, 246, 247 Myrtus communis L . (Myrtaceae), 147,158 M. nivellei Battand., 218, 222 Mystroxylum aethiopicum (Thunb.) Loes.: see Cassine aethiopica Najas (Najadaceae), 265 Narcissus (Amaryllidaceae), 158 Nardurus cynosuroides (Desf.) Trabut (Gramineae), 230 Nardus stricto L . (Gramineae), 155 Nastus borbonicus J. F . Gmelin (Gramineae), 258 Nauclea diderrichii (De Wild. & T h . Durand) Merr. (Rubiaceae), 77 N. latifolia Smith, 83, 85, 105, 107,108 N. pobeguinii (Pobéguin ex Pellegr.) Petit, 83, 91 Neoboutonia macrocalyx Pax (Euphorbiaceae), 181 N. mannii Benth., 253 Neocentema (Amaranthaceae), 111 Neodypsis (Palmaceae), 232, 237 Neophloga (Palmaceae), 232, 235 Nepenthes pervillei Blume (Nepenthaceae), 257 Nephrosperma (Palmaceae), 257 Neptunio olerácea Lour. (Leguminosae: Mimosoideae), 206 Nerium oleander L . (Apocynaceae), 147, 219, 222 Nesiota (Rhamnaceae), 254 N. elliptica (Roxb.) Hook.f., 254 Nesogordonia papaverifera (A. Chev.) Capuron (Steruliaceae), 79,178 N. parvifolia ( M . B . Moss) Capuron, 188 Nestlera (Compositae), 140 Neuracanthus (Acanthaceae), 111
345
Neurada (Neuradaceae), 218 N, procumbens L., 220 Neurotheca congolana D e Wild. & T h . Durand (Gentianaceae), 84 Newbouldia laevis (P. Beauv.) Seemann ex Bureau (Bignoniaceae), 81 Newtonia aubrevillei (Pellegr.) Keay (Leguminosae: M i m o soideae), 82 N. buchananii (Baker) Gilbert & Boutique, 80, 85, 91, 167, 181, 186 N. erlangeri (Harms) Brenan, 188 N. hildebrandtii (Vatke) Torre, 90, 91, 117, 200 N. paucijuga (Harms) Brenan, 187, 188 Nicotiana glauca G r a h a m (Solanaceae), 252 Nirarathamnos (Umbelliferae), 111 Nitella (Characeae), 265 Nitraria refusa (Forssk.) Asch. (Zygophyllaceae), 223, 224, 230 Northea seychellana Hook.f. (Sapotaceae), 257 Nostoc commune Vaucher (Nostocaceae), 53 Notelaea azorica Tutin: see Picconia azorica N. excelsa (Aiton) W e b b & Berth.: see P. excelsa Notholaena (Sinopteridaceae), 238 Notonia (Compositae), 242 Nucularia (Chenopodiaceae), 218 N. perrinnii Battand., 223 Nuxia (Loganiaceae), 237 N. congesta R . Br. L ex Fresen., 74, 122, 165, 167, 176, 196, 199,253 N. floribunda Benth., 165 N. pseudodentata Gilg, 255 N. verticillata L a m . , 258 Nymania (Meliaceae), 137 N. capensis (Thunb.) Lindb., 140,191 Nymphaea (Nymphaeaceae), 265 N. caerulea Savigny, 265 N. lotus L . , 265 Nymphoides ezannoi Berhaut (Menyanthaceae), 204 N. indica (L.) Kuntze, 265 Ochlandra (Gramineae), 237 O. capitata C a m u s , 235, 236 Ochna (Ochnaceae), 83 O. afzelii R . Br. ex Oliver, 105 O. ciliata L a m . , 259 O. holstii Engl., 164 O. leptoclada Oliver, 174 O. manikensis D e Wild., 173 O. membranácea Oliver, 82 O. ovata F . Hoffm., 176 O. pulchra Hook., 96, 97,193, 196 O. schweinfurthiana F . Hoffm., 99,105, 109,174 O. thomasiana Engl. & Gilg, 188 Ochradenus (Resedaceae), 218 O. baccatus Del., 116 Ochrocarpos: see Mammea Ochthocosmus lemaireanus D e Wild. & T h . Durand (Ixonanthaceae), 97 Ocimum (Labiatae), 122 O. suave Willd., 122 Ocotea (Lauraceae), 235, 237 O. borbónica auct.: see O. obtusata O. búllala (Burchell) Bâillon, 165, 246 O. comoriensis Kosterm., 255 O. foetens (Ait.) Benth. & Hook.f., 246, 247, 249 O. gabonensis R . Fouilloy, 83, 246
346
Index ofplant names
Ocotea (Lauraceae)—contd O. oxyphyllum (Harms) J. Léonard, 77 O. kenyensis (Chiov.) Robyns & R . Wilczek, 122,165, 246 Oxytenanthera abyssinica (A. Rich.) M u n r o (Gramineae), 55 O. michelsonii Robyns & R . Wilczek, 85 Ozoroa crassinervia (Engl.) R . & A . Fernandes: see Heeria O. obtusata (Nees) Kostermans, 258 crassinervia O. usambarensis Engl., 85, 164, 187 O. reticulata (Baker f.) R . & A . Fernandes: see H. reticulata Odyssea jaegeri (Pilger) Robyns & Tournay: see Psilolemma jaegeri Pachycarpus lineolatus (Decne.) Bullock (Asclepiadaceae), 100 O. paucinervis (Nees) Stapf, 144, 267 Pachyelasma (Leguminosae: Caesalpinioideae), 74 Oldenburgia arbúsculo D C . (Compositae), 134 Pachypodium (Apocynaceae), 238 Oldfieldia africana Benth. & Hook.f. (Euphorbiaceae), 77 P. geayi Costantin & Bois, 242 O. dactylophylla (Welw. ex Oliver) J. Léonard, 97, 173 P. lamerei Drake, 242 O. somalensis (Chiov.) Milne-Redh., 188 P. lealii W e l w . , 140 Olea (Oleaceae), 130, 226, 227, 228, 255 P. namaquanum (Wyley ex Harv.) W e l w . , 140 O. africana Miller, 115, 121, 122, 129, 135, 167, 182, 193, P. succulentum (L.f.) A . D C , 195 194,195,201,224 Pachystela brevipes (Baker) Bâillon ex Engl. (Sapotaceae), 105, O. capensis L . , 62, 105, 109, 122, 134, 164, 253 109, 172, 188 ss. macrocarpa ( C . H . Wright) I. Verdoorn, 200 P.msolo (Engl.) Engl., 187 O. europaea L„ 149, 152, 154, 157, 158, 159, 226, 227, 228, Paeonia atlántica Kralik ex Trabut (Paeoniaceae), 157 229,251 Pandanus (Pandanaceae), 188, 236, 243, 257 O. foveolata E . Meyer: see Chionanthus foveolatus P. alpestris Martius, 238 O. hochstetteri Baker: see O. capensis P. candelabrum P. Beauv., 83, 262 O. laperrinei Battand. & Trabut, 54, 204, 207, 208, 211, 222 P. goetzei W a r b . , 189 O. woodiana Knobl., 200 P. heterocarpus Balf.f., 258 Oleandro articulata Presl (Oleandraceae), 235 P. hornei Balf.f., 257 Olinia (Oliniaceae), 135,196 Pandiaka carsonii (Baker) C . B . Clarke (Amaranthaceae), 100 O. emarginata Burtt Davy, 195 Panicum aldabrense Renvoize (Gramineae), 259 Oncinoíis inhandensis J. M . W o o d & Evans (Apocynaceae), 200 P. baumannii K . Schum., 174 Oncostemum (Myrsinaceae), 232, 237 P. coloratum L . , 116, 126, 194 Onobrychis argéntea Boiss. (Leguminosae: Papilionoideae), 230 P. deustum Thunb., 182 Ononis (Leguminosae: Papilionoideae), 147 P. dregeanum Nees, 239 O. atlántica ha\l, 158 P. fulgens Stapf: see P. baumannii O. polysperma Barr. & Murbeck, 227 P. griffonii Franchet, 84 Ophiobotrys (Flacourtiaceae), 74 P. heterostachyum Hackel, 90, 97 Ophrys (Orchidaceae), 158 P. kalaharense M e z , 193 Opilia celtidifolia (Guill. & Perr.) Endl. ex Walp. (Opiliaceae), P. laetum Kunth, 204 109 P. lanipes M e z , 193 Oplismenus hirtellus (L.) P . Beauv. (Gramineae), 90 P. lindleyanum Nees ex Steud., 84 Opuntia (Cactaceae), 159, 231, 254 P. luridum Hackel, 239 Orbea (Asclepiadaceae), 113 P. maximum Jacq., 85,101, 236, 243, 252 Orchis (Orchidaceae), 158 P. natalense Hochst., 193 Oreobambos buchwaldii K . Schum. (Gramineae), 55 P. parvifolium L a m . , 84, 266 Oricia bachmannii (Engl.) I. Verdoorn (Rutaceae), 199 P. phragmitoides Stapf, 85, 174 Ormenis multicaulis Braun-Blanquet ex Maire (Compositae), P. pilgeri M e z , 84 152 P. pusillum Hooi.1, 211 Ornithogalum (Liliaceae), 158 P. repens L . , 55,100 Orothamnus zeyheri Pappe (Proteaceae), 135 P. subalbidum Kunth, 209, 266 Oryza longistaminata Chev. & Roehr. (Gramineae), 100,108, 265 P. turgidum Forssk., 115, 204, 207, 212, 213, 216, 218, 219, O. perennis auct.: see O. longistaminata 220, 221, 224 Oryzopsis caerulescens (Desf.) Hackel (Gramineae), 222 Pappea capensis Eckl. & Zeyh. (Sapindaceae), 98, 117, 121, 129, Osmunda regalis L . (Osmundaceae), 156 137,140,141,189,201 Osteospermum (Compositae), 141, 194 Paramacrolobium coeruleum (Taubert) J. Léonard (Leguminosae: Caesalpinioideae), 74,188 O. scariosum D C , 194 Parinari capensis Harv. (Chrysobalanaceae), 101, 173, 201 Ostryoderris stuhlmannii (Taubert) Harms: see Xeroderris stuhlmannii P. congensis F . Didr., 83, 176 Osyris sp. (Santalaceae), 129, 135, 153, 154, 194, 195 P. congolona T h . & H . Durand, 83 Othonna protecta Dinter (Compositae), 142 P. curatellifolia Planchón ex Benth., 85, 87, 93, 96, 97, 98, Otoptera (Leguminosae: Papilionoideae), 243 99,105, 129, 173, 174, 189 Olostegia (Labiatae), 111 P. excelsa Sabine, 71, 74, 81, 82, 85, 90, 92, 164, 176, 178, Ottelia ulvifolia (Planchón) Walp. (Hydrocharitaceae), 265 181 Oubanguia africana Bâillon (Scytopetalaceae), 83 P. glabra Oliver: see Maranthes glabra Ouratea (Ochnaceae), 74, 83 P. goetzeniana Engl.: see M . goetzeniana Oxalis (Oxalidaceae), 141, 194 P. polyandra Benth.: see M . polyandra O. depressa Eckl. & Zeyh., 194 Parkia bicolor A . Chev. (Leguminosae: Mimosoideae), 55, 77, Oxystigma (Leguminosae: Caesalpinioideae), 74 82 O. mannii (Bâillon) Harms, 83 P. biglobosa (Jacq.) Benth., 83, 85,105
Index of plant names
P. clappertoniana Keay: see P. biglobosa P. filicoidea W e l w . ex Oliver, 74, 91, 171, 172, 181, 186, 187, 188 Parkinsonia aculeata L. (Leguminosae: Caesalpinioideae), 252 P. africana Sond., 140, 143, 144,191 Parmelia (Parmeliaceae), 142 P. vagans Nylander, 53 Parnassia palustris L . (Parnassiaceae), 155 Paspalidium geminatum (Forssk.) Stapf (Gramineae), 55, 265 Paspalum commersonii L a m . (Gramineae): see P. scrobiculatum P. orbiculare Forster: see P. scrobiculatum P. scrobiculatum L., 100, 202, 266 P. vagin*tum Swartz, 262, 263 Passerina (Thymelaeaceae), 134, 167 P. filiformis L., 134 P. montana Thoday, 195 Paullinia pinnata L . (Sapindaceae), 83 Pavonia urens Cav. (Malvaceae), 122 Peddiea fischeri Engl. (Thymelaeaceae), 74 P. thomensis Engl. & Gilg, 253 Peganum harmala L . (Zygophyllaceae), 227, 266 Pegolettia retrofracta (Thunb.) Kies (Compositae), 195 Pelargonium (Geraniaceae), 139 P. cotyledonis (L.) L'Hérit., 254 P. cristophoranumVetàc, 116 P. otaviense Knuth, 142 P. rôssingense Din ter: see P. otaviense Pellaea (Sinopteridaceae), 99, 238 Peltophorum africanum Sond. (Leguminosae: Caesalpinioideae), 95, 96, 98, 201 Pemphis acidula Forst. (Lythraceae), 259 Pennisetum (Gramineae), 169, 211 P. mezianum Leeke, 126, 127, 128 P. pedicellatum Trin., 209 P. polystachion (L.) Schultes, 84, 107, 252 P. purpureum Schumach., 50, 85 P. ramosum (Hochst.) Schweinf., 209 P. schimperi Steud., 130 P. stramineumA. Peter, 126,127 P. unisetum (Nées) Benth., 107, 174 Pentaclethra macrophylla Benth. (Leguminosae: Mimosoideae), 81,253 Pentadesma lebrunii Staner (Guttiferae), 79, 85 Pentanopsis (Rubiaceae), 111 Pentaschistis (Gramineae), 134, 169 P. humbertii A . C a m u s , 239 P. patula (Nées) Stapf, 135 P. perrieri A . C a m u s , 239 P. pictigluma (Steud.) Pilger, 211 P. tysonii Stapf, 169 Pentzia (Compositae), 137, 139, 193 P. globosa Less., 195 P. incana (Thunb.) Kuntze, 193 P. monodiana Maire, 222 P. sphaerocephala D C , 195 Peperomia (Piperaceae), 165, 236, 237, 253 P. fernandopoiana C . D C , 82 P. staudtii Engl.: see P. fernandopoiana Pergularia daemia (Forssk.) Chiov. (Asclepiadaceae), 114 Pericopsis angolensis (Baker) van M e e u w e n (Leguminosae: Papilionoideae), 93, 95, 96, 98, 174 P. elata (Harms) van M e e u w e n , 77 P. laxiflora (Benth. ex Baker) van M e e u w e n , 85, 105,107 Periploca laevigata Aitón (Asclepiadaceae), 153, 227, 228, 229, 251
347
Perotis patens G a n d . (Gramineae), 243 Persea azorica Seub. (Lauraceae): see Laurus azorica P. indica (L.) Spreng., 246, 247, 249 Petalidium (Acanthaceae), 140 P. angustifolium P. G . Meyer, 142 P. engleranum C. B . Clarke, 267 P. giessii P. G . Meyer, 142 Petersianthus macrocarpus (P. Beauv.) Liben (Lecythidaceae), 77,81,172 Petrobium (Compositae), 254 P. arboreum R . Br., 254 Phaeoptilum (Nyctaginaceae), 137 P. spinosum Radlk., 191, 193 Pharnaceum acidum Hook.f. (Aizoaceae), 254 Phaseolus lunatus L . (Leguminosae: Papilionoideae), 252 P. vulgaris L., 252 Philippia (Ericaceae), 132,167, 168, 189, 236, 237, 238, 258 P. abietina (Willd.) Klotzsch, 258 P. benguelensis (Welw. ex Engl.) Britten, 99 P. chamissonis Klotzsch, 134 P. comorensis Engl., 255 P. mafiensis Engl., 189 P. montana (Willd.) Klotzsch, 258 P. simii S. Moore, 189 P. thomensis Henriq., 253 Phillyrea angustifolia L . (Oleaceae), 149, 152, 153, 155, 158, 159,226,228 P. latifolia L . : see P. angustifolia P. media L . : see P. angustifolia Philoxerus vermicularis (L.) P. Beauv. (Amaranthaceae), 262 Phoenicophorium (Palmaceae), 257 Phoenix atlántica A . Chev. (Palmaceae), 246, 251 P. canariensis Chabaud, 246 P. dactylifera L., 216, 219, 226 P. reclinata Jacq., 83,122, 181, 188, 200, 211 Phormium tenax J. R . & G . Forster (Agavaceae), 254 Phragmites (Gramineae), 55 P. australis (Cav.) Trin. ex Steud., 218, 219, 222, 223, 224, 265 P. mauritianus Kunth, 265 Phyla nodiflora (L.) Greene (Verbenaceae), 145 Phylica (Rhamnaceae), 132, 134 P. buxifolia L., 134 P. leucocephala Cordem.: see P. nítida P. mauritiana Bojer ex Baker.: see P. nítida P. nitida L a m . , 258 P. oleifolia Vent., 134 P. paniculata Willd., 134 P. ramosissima D C . P. villosa Thunb., 134 Phyllanthus comorensis Engl. (Euphorbiaceae), 255 P. discoideus (Bâillon) Muell. Arg., 98, 167, 199 P. maderaspatensis L., 191 P. muelleranus (Kuntze) Exell, 173 P. verrucosus Thunb., 201 Phymaspermum (Compositae), 137 Phymatodes scolopendria (Burm.f.) Ching (Polypodiaceae), 189 Picconia (Oleaceae), 246 P. azorica (Tutin) Knobl., 246, 247 P. excelsa (Aitón) D C , 246, 247 Picralima nitida (Stapf) T h . & H . Durand (Apocynaceae), 83 Piliostigma reticulatum ( D C ) Hochst. (Leguminosae: Caesalpinioideae), 105,107, 108, 206 P. thonningii (Schumach.) Milne-Redh., 83, 85, 87, 95, 96, 105,107,173,174,189,210,214
348
Index of plant names
Pilostyles aethiopica W e l w . (Rafflesiaceae), 93 Pilotrichella (Metioriaceae), 81 Pimpinella villosa Schousboe (Umbelliferae), 157 Pinguicula vulgaris L . (Lentibulariaceae), 155 Pinus canariensis Chr. Smith ex D C . (Pinaceae), 246, 248, 249 P. halepensis Miller, 51, 146, 149, 150, 151, 152, 153, 154, 155,157,160,226,229,231 P. nigra Arnold, 152 P. pinaster Aitón, 135, 146, 149, 150,152, 155 Piper capense L.f. (Piperaceae), 74 Piptadeniastrum africanum (Hook.f.) Brenan (Leguminosae: Mimosoideae), 75, 78, 79, 82, 172, 181 Piptatherum coerulescens (Desf.) Hackel: see Oryzopsis coerulescens Pisonia aculeata L . (Nyctaginaceae), 200 Pistacia (Anacardiaceae), 227 P. atlántica Desf., 147, 149,150, 157, 158, 222, 226, 227 P. lentiscus L . , 115, 121, 149, 151, 152, 153, 154, 157, 158, 159,226,227,228 P. terebinthus L., 149 Pistia stratiotes L . (Araceae), 265 Pittosporum (Pittosporaceae), 238 P. coriaceum Dryander ex Aiton, 246, 247 P. lanceolatum Cordem.: see P. senada P. senada Putterl, 258 P. viridiflorum Sims, 62, 74, 195, 196 Pituranthos battandieri Maire (Umbelliferae), 221 Placopoda (Rubiaceae), 111 Pladaroxylon (Compositae), 254 P. leucadendron (Forster f.) Hook.f., 254 Plagiochila (Plagiochilaceae), 81 Plagiochloa (Gramineae), 134 Plantago (Plantaginaceae), 236, 246 P. ciliata Desf., 220 P. coronopus L., 266 P. robusta Roxb., 254 Platycelyphium (Leguminosae: Papilionoideae), 111 P. voense (Engl.) Wild, 114 Platycerium (Polypodiaceae), 235 P. elephantotis Schweinf., 182 Platypterocarpus (Celastraceae), 162 Plectranthus (Labiatae), 241 P. ignarius (Schweinf.) A g n e w , 120 Pleiomeris (Myrsinaceae), 246 P. canariensis (Willd.) A . D C , 246 Pleuropterantha (Amaranthaceae), 111 Pleurostylia africana Loes. (Celastraceae), 188 Plinthus (Aizoaceae), 140, 193 Plumbago auriculata L a m . (Plumbaginaceae), 201 P. capensis Thunb.: see P. auriculata P. zeylanica L., 90 Poa (Gramineae), 169 P. ankaratrensis A. C a m u s , 239 P. madecassa A. C a m u s , 239 Podalyria (Leguminosae: Papilionoideae), 132 Podocarpus (Podocarpaceae), 165, 237 P. elongatus (Aiton) L'Hérit. ex Pers., 135 P. ensiculus Melville: see P. henkelii P. fakatus (Thunb.) R . Br. ex Mirbel, 165,181, 199, 200 P. gracilior Pilger: see P. falcatus P. henkelii Stapf, 165 P. latifolius (Thunb.) R . Br. ex Mirbel, 164, 165, 167, 181, 195,196, 199 P. madagascariensis Baker, 236 P. mannii Hook.f., 253
P. milanjianus Rendle: see P. latifolius P. rostratus Laurent, 238 P. usambarensis Pilger var. dawei (Stapf) Melville: see P. falcatus Poga (Rhizophoraceae), 74 Pogonarthria squarrosa (Licht. ex R o e m . & Schult.) Pilger (Gramineae), 191,193,194, 243 Polycarpaea fragilis Del. (Caryophyllaceae): see P. repens P. repens (Forssk.) Asch. & Schweinf., 220 Polyceratocarpus scheffleri Engl. & Diels (Annonaceae), 187 Polygala (Polygalaceae), 132 P. arenaria Willd., 178 P. balansae Cosson, 154 P. myrtifolia L., 134 Polygonum (Polygonaceae), 266 P. acuminatum Kunth, 266 P. pulchrum Blume, 265 P. salicifolium Brouss. ex Willd., 265 P. strigosum R . Br., 265 Polypogon monspeliensis (L.) Desf. (Gramineae), 246 Polyscias fulva (Hiern) H a r m s (Araliaceae), 74, 211 P. quintasii Exell, 253 Polysphaeria multiflora Hiern (Rubiaceae), 259 Popowia (Annonaceae), 98 P. obovata (Benth.) Engl. & Diels, 90, 98 Populus alba L . (Salicaceae), 150 P. euphratica Oliver, 219 P. ilicifolia (Engl.) Rouleau, 117 P. trémula L . , 149,156 Portulaca olerácea L . (Portulacaceae), 254 Portulacaria afra Jacq. (Portulacaceae), 137,140, 141, 201 Poskea (Globulariaceae), 111, 147 Potamogetón richardii Solms-Laub. (Potamogetonaceae), 265 P. schweinfurthii A. Bennett, 265 Poterium spinosum L . (Rosaceae), 159 Premna hildebrandtii Giirke (Verbenaceae), 114 P. quadrifolia Schumach. & Thonn., 178 P. resinosa (Hochst.) Schauer, 114 Primula vulgaris Hudson (Primulaceae), 155 Prosopis africana (Guill. & Perr.) Taubert (Leguminosae: Mimosoideae), 105, 107, 209 Protarum (Araceae), 257 Protea (Proteaceae), 93, 99, 132, 134 P. arbórea Houtt., 134 P. caffra Meissner, 96 P. glabra Thunb., 134 P. laurifolia Thunb., 134 P. longiflora L a m . , 134 P. lorifolia (Salisb. ex Knight) Fourc, 134 P. madiensis Oliver, 105 P. nereifoliaR. Br., 134 P. obtusifolia Buek, 134 P. petiolaris W e l w . ex Engl., 173 P. repens (L.)L., 134, 135 P. susannae E . P . Phillips, 134 Protorhus (Anacardiaceae), 199, 236 P. buxifolia H . Perrier, 237 P. deflexa H . Perrier, 241 P. humbertii H . Perrier, 241 P. longifolia (Bernh.) Engl., 199, 201 P. perrieri Courchet, 241 Prunus africana (Hook.f.) Kalkman (Rosaceae), 80, 122, 164, 165,167, 181,253,255 P. avium L., 149,152,157 P. lusitanica L., 149
Index of plant names
P. padus L . , 147,149, 156 P. pérsica (L.) Batsch, 252 P. prostrata Labill., 156, 158 Pseudagrostistachys africana (Muell. Arg.) Pax & Hoffm. (Euphorbiaceae), 253 Pseudocedrela (Meliaceae), 103 P. kotschyi (Schweinf.) Harms, 83, 85, 105, 107,108, 209 Pseudolachnostylis (Euphorbiaceae), 87 P. maprouneifolia Pax, 31, 95, 96, 97 Pseudoprosopis fischeri (Taubert) H a r m s (Leguminosae: M i m o soideae), 97 Pseudosalacia (Celastraceae), 199 Pseudospondias microcarpa (A. Rich.) Engl. (Anacardiaceae), 181,253 Psiadia (Compositae), 237, 238, 258 P. altissima ( D C . ) Benth. & Hook.f., 236 P. arabica Jaub. & Spach: see P. punctulata P. punctulata ( D C . ) Vatke, 115 P. schweinfurthii Balf.f., 255 Psidium cattleianum Sabine (Myrtaceae), 236, 257 P. guajava L . , 83, 236, 252, 254 Psilocaulon salicornioides (Pax) Schwantes (Aizoaceae), 142 Psilolemmajaegeri (Pilger) S. M . Phillips (Gramineae), 267 Psilonema (Cruciferae), 111 Psilotrichum (Amaranthaceae), 111 Psoralea obtusifolia D C . (Leguminosae: Papilionoideae), 144 P.pinnataL., 134 P. plicata Del., 219 Psorospermum febrifugum Spach (Guttiferae), 85,174 Psychotria capensis (Eckl.) Vatke (Rubiaceae), 200 P. peduncularis (Salisb.) Steyerm., 91 Ptaeroxylon obliquum (Thunb.) Radlk. (Ptaeroxylaceae), 96, 165, 200 Pteleopsis anisoptera (Welw. ex Lawson) Engl. & Diels (Combretaceae), 98 P. díptera (Welw.) Engl. & Diels, 80, 172 P. myrtifolia (Lawson) Engl. & Diels, 200 P. suberosa Engl. & Diels, 107 Pteridium aquilinum (L.) K u h n (Pteridiaceae), 96, 152, 157, 236 Pteris (Pteridiaceae), 75 P. vittata L . , 252 Pterocarpus (Leguminosae: Papilionoideae), 98 P. angolensis D C , 93, 95, 96, 97, 98,173, 174 P. antunesii (Taubert) Harms, 90 P. erinaceus Poir., 83, 85, 105 P. lucens Guill. & Perr., 209, 211 P. mildbraedii Harms, 74 ss. usambarensis (Verde.) Polhill, 187 P. rotundifolius (Sond.) Druce, 95, 96, 98, 99, 201 P. santalinoides L'Hérit. ex D C , 176 Pterocelastrus (Celastraceae), 96, 196 P. tricuspidatus Sond., 135 Pterodiscus (Pedaliaceae), 140 Pterolobium stellatum (Forssk.) Brenan (Leguminosae: Caesalpinioideae), 115 Pteronia (Compositae), 135,137 P. glauca Thunb., 140 Pterygota macrocarpa K . Schum. (Sterculiaceae), 79, 81 Ptilotrichum spinosum (L.) Boiss.: see Alyssum spinosum Puccionia (Cruciferae), 111 Púnica granatum L . (Punicaceae), 252 Pupalia lappacea (L.) Juss. (Amaranthaceae), 90 Putterlickia pyracantha (L.) Szyszyl. (Celastraceae), 135 Pycnanthus angolensis (Welw.) W a r b . (Myristicaceae), 80, 81, 85, 172, 181
349
Pycnocoma littoralis Pax (Euphorbiaceae), 189 Pycreus aethiops (Welw. ex Ridley) C . B . Clarke (Cyperaceae), 100 Pygeum africanum Hook.f.: see Prunus africana Pyrenacantha malvifolia Engl. (Icacinaceae), 114 Pyrus cossonii Rehder (Rosaceae), 149 P. gharbiana Trabut, 149 P. longipes Coss. & Durieu: see P. cossonii P. mamorensis Trabut, 149,151 Quercus afares Pomel (fa*gaceae), 146, 149, 151,156, 157 Q. calliprinos W e b b : see Q. coccifera Q. coccifera L . , 146, 147, 149, 150,152, 154,158,159, 226 Q.fa*ginea L a m . , 146, 149,150, 151,152,155, 156, 157 Q. ilex L . , 146, 147, 149, 150, 151, 152, 153, 154, 156, 157, 226,229,231 Q. lusitanica n o m . ambig.: see Q. fa*ginea Q. pyrenaicaWiWA., 146, 149, 150, 156, 157 Q. suberL., 48, 146, 147, 149, 150, 151, 152, 154, 155, 157, 237 Q. toza Bast.: see Q. pyrenaica Quivisia oppositifolia Cav.: see Turraea oppositifolia Racopilum speluncae P. Beauv. (Racopilaceae), 182 Ramalina (Ramalinaceae), 224 Randonia africana Coss. (Resedaceae), 221 Ranunculus (Ranunculaceae), 236 Rapanea melanophloeos (L.) M e z (Myrsinaceae), 122, 135, 165, 167,196, 200 Raphia (Palmaceae), 83, 91, 188 R. australis Oberm. & Strey, 200 R. farinifera (Gaertner) Hylander, 181 Rauvolfia caffra Sond. (Apocynaceae), 187, 200 R. nana E . A . Bruce, 173 R. vomitoria Afzel., 80, 253 Ravenala (Strelitziaceae), 232 R. madagascariensis Adans., 235, 236, 257 Ravensara (Lauraceae), 232, 235, 241 Reaumuria hirtella Jaub. & Spach (Tamaricaceae), 221 R. muricata Jaub. & Spach: see R. vermiculata R. vermiculata L . , 230 Redfieldia hitchco*ckii A . C a m u s (Gramineae), 238 Rendlia altera (Rendle) Chiov. (Gramineae), 196 Reseda battandieri Pitard (Resedaceae), 227 R. villosa Coss., 221 Restio (Restionaceae), 132 Retama bovei Spach (Leguminosae: Papilionoideae): see R. monosperma R. monosperma (L.) Boiss., 152, 231 R. retam W e b b , 147, 220, 224 Rhamnus alaternus L . (Rhamnaceae), 153 R. alpinusL., 158 R. catharticus L . , 151 R. frángula L . , 156 R. latifolia L'Hérit., 247 R. oleoides L . , 153,158, 227 R. prinoides L'Hérit., 195 R. staddo A . Rich., 121 Rhanterium (Compositae), 218 Rhigozum angolense B a m p s (Bignoniaceae), 145 R. brevispinosum Kuntze, 95, 193 R. madagascariensis Drake, 242 R. obovatum Burchell, 96,140, 193, 195 R. trichotomum Burchell, 139,140, 191, 193 R. virgatum M e r x m . & A . Schreiber, 95,140
350
Index ofplant names
Rhipsalis (Cactaceae), 165, 174, 236 Rhizophora (Rhizophoraceae), 55, 261, 262, 263, 264 R. harrisonii Leechman, 253, 261, 262 R. mangle L . , 261, 262 R. mucronata L a m . , 259, 261, 263, 264 R. racemosa G . F . W . Meyer, 261, 262 Rhodognaphalon schumannianum A . Robyns (Bombacaceae), 188 Rhoicissus digitata (L.f.) Gilg & Brandt (Vitaceae), 201 R. tomentosa (Lam.) Wild & R . B . D r u m m o n d , 200 R. tridentata (L.f.) W ü d & R . B . D r u m m o n d , 98, 201 Rhus (Anacardiaceae), 96, 201 R. albida Schousboe, 251 R. chirindensis Baker f., 96 R. ciliata Licht. ex Schultes, 193, 194,195 R. crenata Thunb., 135 R. dregeana Sond., 193 R. erosa Thunb., 193, 194, 195 R. glauca Thunb., 135 R. incana Mill., 222 R. laevigata L . , 135 R. lancea L.f., 141,144, 193, 194, 195 R. leptodictya Diels, 96 R. longipes Engl., 109 R. lucida L . , 135 R. marlothii Engl., 193 R. mucronata Thunb., 135 R. natalensis Bernh. ex Krause, 109,115, 129, 182 R. oxyacantha Shousb., 227 R. pentaphylla (Jacq.) Desf., 149,158, 227, 228 R. pyroides Burchell, 193, 195 R. quartiniana A. Rich., 91 R. somalensis Engl., 115 R. taratana (Baker) H . Perrier, 237 R. thyrsiflora Balf.f., 255 R. tomentosa L . , 135 R. tripartita (Ucria) Grande, 222 R. undulata Jacq., 140, 141, 193, 195 R. vulgaris Meikle, 121, 207 Rhynchelytrum amethysteum (Franchet) Chiov. (Gramineae), 101, 173 R. repens (Willd.) C . E . Hubbard, 101, 145, 252 R. villosum (Pari, ex Hook.f.) Chiov.: see R. repens Rhynchocalyx (Lythraceae), 199 Rhynchosia (Leguminosae: Papilionoideae), 220 R. candida (Welw. ex Hiern) Torre, 144 R. memnonia (Del.) Boiss., 222 R. totta Thunb., 194 Rhynchospora candida (Nées) Boeck (Cyperaceae), 84 R. corymbosa (L.) Britten, 31, 84 R. holoschoenoides (L. C . Rich.) Herter, 84 R. rubra (Lour.) Makino, 84 R. rugosa (Vahl) Gale, 84 Rhytachne rottboellioides Desv. (Gramineae), 32, 84, 108 Ribes alpinum L . (Grossulariaceae), 158 R. uva-crispa L . , 158 Ricinodendron heudelotii (Bâillon) Pierre ex Pax (Euphorbiaceae), 74, 78,79, 81,172,186,187,188 R. rautanenii Schinz, 90, 97 Ricinus communis L . (Euphorbiaceae), 252 Rinorea (Violaceae), 74, 235 R. malembaensis T a ton, 172 Riseleya griffithii Hemsley (Euphorbiaceae), 257 Romulea (Iridaceae), 158 Rosa (Rosaceae), 151, 157
Roscheria (Palmaceae), 257 R. melanochaetes (H. A . Wendl.) H . A . Wendl. ex Balf.f., 257 Rosmarinus eriocalix Jordan & Fourr. (Labiatae), 155, 226, 229 R. officinalis L . , 154, 226 R. toumefourtii D e N o é ex Turrill: see R. eriocalix Roíala pterocalyx A. Raynal (Lythraceae), 204 Rothia (Leguminosae: Papilionoideae), 243 Rubus (Rosaceae), 151, 167, 236, 254 R. pinnatus Willd., 253 R. ulmifolius Schott, 151,157 Ruellia (Acanthaceae), 140 R. insignis Balf.f., 115, 255 Ruschia (Aizoaceae), 139, 140 R. unidens Schwantes, 195 Ruscus aculeatus L . (Ruscaceae), 151, 157 Ruta (Rutaceae), 228 Rytigynia umbellulata (Hiern) Robyns (Rubiaceae), 98 Sacciolepis africana C . E . Hubbard & Snowden (Gramineae), 100 Sacoglottis gabonensis (Bâillon) Urban (Houmiriaceae), 76, 174 Salacia (Celastraceae), 75, 241 S. kraussii (Harv.) Harv., 202 Salicornia (Chenopodiaceae), 267 S. arabica L . , 223, 230 S. fruticosa L . : see S. arabica Salix alba L . (Salicaceae), 150 S. cinérea L . , 151,156 S. purpurea L . , 156 Salsola (Chenopodiaceae), 144, 193, 223, 224 S. aphylla L.f., 139, 142 S. baryosma (Schultes) D a n d y , 223 S. foetida Del. ex Sprengel: see S. baryosma S. longifolia Forssk., 230, 266 S. nollothensis Aellen, 142 S. oppositifolia Desf.: see S. longifolia S. sieberi Presl, 223, 229 S. tetrágono Del., 223, 229, 230 S. tuberculata (Moq.) Schinz, 139, 267 S. vermiculata L . , 228, 229, 230, 266 S. zeyheri ( M o q . ) Schinz, 144 Salvadora pérsica L . (Salvadoraceae), 114, 119, 121, 128, 144, 207, 219, 220, 224, 267 Salvia (Labiatae), 221 S. aegyptiaca L . , 221 Salvinia auriculata Aublet (Salviniaceae), 265 S. molesta Mitchell, 265, 266 Sambucus africana Standley (Caprifoliaceae), 167 Samolus valerandi L . (Primulaceae), 145 Sanguisorba (Rosaceae), 246 Sanícula (Umbelliferae), 236 S. europaea L . , 156 Sansevieria (Agavaceae), 98, 114,115, 116,129, 188, 201 S. arborescens G é r o m e & Labroy, 114 S. cylindrica Bojer, 145, 174 S. ehrenbergii Schweinf. ex Baker, 117,128 S. liberica G é r o m e & Labroy, 178 Santiria trimera (Oliver) Aubrév. (Burseraceae), 82, 83 Sapium bussei Pax: see Excoecaria bussei S. ellipticum (Hochst.) Pax, 74, 199, 253 Sarcocaulon (Geraniaceae), 137, 139 S. marlothii Engl.: see S. mossamedense S. mossamedense (Welw. ex Oliver) Hiern, 142, 144, 145 S. spinosum (Burm.f.) Kuntze, 142
Index of plant names
Sarcolaena oblongifolia Gérard (Sarcolaenaceae), 237 Sarcophtynium (Marantaceae), 75 Sarcopoterium spinosum (L.) Spach: see Poterium spinosum Sarcostemma (Asclepiadaceae), 116 5. daltonii Decne., 251 S. viminale (L.) Aiton f„ 114, 115,117, 191,201 Saxymolobium holubii (Scott Elliot) Bullock (Asclepiadaceae), 100 Scabiosa columbaria L . (Dipsaceae), 194 Scaevola sp. (Goodeniaceae), 257, 258 Schefflera (Araliaceae), 236, 237 S. barteri (Seemann) H a r m s , 82, 83 S. bojeri R . Viguier, 237, 238 S. mannii (Hook.f.) H a r m s , 253 S. umbellifera (Sond.) Bâillon, 165 Schefflerodendron usambarense H a r m s (Leguminosae: Papilionoideae), 74, 187 Schismus barbatus Juel (Gramineae), 50 Schizachyrium brevifolium (Swartz) Nées ex Büse (Gramineae), 174 5. exile (Hochst.) Pilger, 210 S. platyphyllum Stapf, 50 S. sanguineum (Retz.) Alston, 85,101 S. semiberbe Nées: see S. sanguineum S. thollonii Stapf, 173 Schmidtia kalahariensis Stent (Gramineae), 145, 191,193 S. pappophoroides J. A . Schmidt, 114, 137, 145, 193, 206, 252 Schoenefeldia gracilis Kunth (Gramineae), 108, 176, 206, 207, 208, 212, 213 Schotia (Leguminosae: Caesalpinioideae), 201 S. afra (L.) Thunb., 137,140, 201 S. africana (Bâillon) Keay, 77 S. brachypetala Sond., 96, 200 S. latifolia Jacq., 137, 200, 201 Schouwia (Cruciferae), 218 Schrebera alata (Hochst.) W e l w . (Oleaceae), 115, 129, 167 S. arbórea A. Chev., 176 S. trichoclada W e l w . , 174 Schumanniophyton problematicum (A. Chev.) Aubrév. (Rubiaceae), 75 Scilla (Liliaceae), 158 S. nervosa (Burchell) Jessop, 194 Scirpus cubensis Poeppig & Kunth (Cyperaceae), 266 S. holoschoenus L . , 218, 219, 222, 266 5. inclinatus (Del.) Asch. & Graebner, 265 S. littoralis Schrader, 144 S. microcephalus (Steud.) Dandy, 100 Scleria aterrima (Ridley) Napper (Cyperaceae), 84 S. bulbifera Hochst. ex A . Rich., 100 S. nutans Kunth, 266 S. nyasensis C . B . Clarke, 266 Sclerocarya (Anacardiaceae), 242 S. birrea (A. Rich.) Hochst., 105, 107, 116, 129, 209, 210, 212,213 S. caffra Sond., 95, 96, 99,174, 189, 201, 243 Sclerocephalus arabicus Boiss. (Caryophyllaceae), 252 Sclerodactylon macrostachyum (Benth.) A . C a m u s (Gramineae), 259 Sclerosciadium nodiflorum Bail (Umbelliferae), 229 Scolopia mundii (Eckl. & Zeyh.) W a r b . (Flacourtiaceae), 165, 195, 196, 199 Scorodophloeus fischeri (Taubert) J. Léonard (Leguminosae: Caesalpinioideae), 117,188 S. zenkeri H a r m s , 77
351
Scutia myrtina (Burm.f.) Kurz (Rhamnaceae), 115, 121, 182, 201 Scytopetalum pierreanum (De Wild.) V a n Tiegh. (Scytopetalaceae), 83 Securidaca longepedunculata Fres. (Polygalaceae), 85, 174, 189, 210,211 Securinega virosa (Roxb. ex Willd.) Bâillon (Euphorbiaceae), 98, 178, 222 S. seyrigii Leandri, 241 Seddera latifolia Hochst. & Steud. (Convolvulaceae), 113 Sedum madagascariense H . Perrier (Crassulaceae), 239 Seetzenia africana R . Br. (Zygophyllaceae), 219 S. orientalis Decne.: see S. africana Sehima ischaemoides Forssk. (Gramineae), 108, 213 Selaginella (Selaginellaceae), 238, 242 S. echinata Baker, 238 5. scandens (P. Beauv.) Spring, 84 Selago (Scrophulariaceae), 135,140 Senecio (Compositae), 53,139,141, 237, 238, 242 subgen. Dendrosenecio, 169 S. anteuphorbium (L.) Hook.f., 224, 227, 228, 229 S. bojeri ( D C . ) Robyns, 182 S. coronatus (Thunb.) Harv., 194 S. erubescens Aiton, 194 S. leucadendron (Forster f.) Hemsley: see Pladaroxylon leucadendron S. longiflorus ( D C . ) Schultz-Bip., 142 S. petitianus A . Rich., 115 5. prenanthiflorus ( D C . ) Hemsley: see Lachanodes arbórea S. redivivus Mabberley: see Lachanodes arbórea S. stuhlmannii Klatt, 182 Sericocomopsis (Amaranthaceae), 111 5. hildebrandtii Schinz, 114,120 S. pallida (C. B . Clarke) Schinz, 114 Serruria (Proteaceae), 132 Sesamothamnus (Pedaliaceae), 113 S. benguellensis W e l w . , 140 S. guerichii (Engl.) E . A . Bruce, 140 S. lugardii N . E . Br., 96 S. n'vaeEngl., 114 Sesbania sesban (L.) Merrill (Leguminosae: Papilionoideae), 266, 267 Sesuvium (Aizoaceae), 144 S. digynum W e l w . ex Oliver: see S. sesuvioides S. portulacastrum L . , 145, 262, 263 S. sesuvioides (Fenzl) Verde, 142 Setaria (Gramineae), 169,189 S. anceps Stapf ex Massey: see S. sphacelata S. chevalieri Stapf, 83, 129 S.flabellata Stapf, 194 S. holstii Herrm.: see S. incrassata S. hom*onyma (Steud.) Chiov., 90 S. incrassata (Hochst.) Hackel, 108, 116 S. lynesii Stapf & C . E . Hubbard, 209 S. nigrirostris (Nées) T h . Durand & Schinz, 194 S. pallide-fusca (Schumach.) Stapf & C . E . Hubbard, 209, 210 S. sphacelata (Schumach.) Stapf & C . E . Hubbard ex M . B . M o s s , 84,100,107,122,194,239 S. verticillata (L.) P. Beauv., 254 Sideroxylon (Sapotaceae), 251, 257 S. bojeranum D C : see S. cinereum S. buxifolium Hutch., 115 S. cinereum L a m . , 257 S. collinum Lecomte, 241 S. galeatum ( A . W . Hill) Baehni, 258
352
Index ofplant names
Sideroxylon (Sapotaceae)—contd S. inerme L . , 135, 188, 189, 200, 201, 259 S. majus (Gaertner f.) Baehni, 258 S. marmulano Banks ex Lowe, 246, 247, 251 Sieglingia decumbens (L.) Bernh. (Gramineae), 155 Silène (Caryophyllaceae), 147 Simocheilus (Ericaceae), 132 Sinapidendron (Cruciferae), 246 Sindoropsis (Leguminosae: Caesalpinioideae), 74 Sisyndite (Zygophyllaceae), 137 Sium helenianum Hook.f. (Umbelliferae), 254 Smilax áspera L . (Smilacaceae), 151, 153 S. kraussiana Meissner, 96 Smithia elliotii Baker f. (Leguminosae: Papilionoideae), 266 Socotora (Asclepiadaceae), 111 S. visciformis (Vatke) Bullock, 113 Socotranthus (Asclepiadaceae), 111 Solanum (Solanaceae), 254 S. albicaule Kotschy ex Dunal, 204 S. auriculatum Aiton, 236 S. dubium Fresen., 212 S. incanum L . , 122 S. indicum L . ss. grandifrons Bitter, 122 Solidago sempervirens L . (Compositae), 246 S. virgaurea L . , 155 Sonchus (Compositae), 246 5. chevalieri (O. Hoffm. & Muschler) Dandy: see Launea chevalieri S. daltonii W e b b , 252 S. nanus Sond. ex Harv., 194 5. pinnatifidus Cav., 226, 228 Sonneratia alba Smith (Sonneratiaceae), 259, 261, 263 Sorbus aria (L.) Crantz (Rosaceae), 149, 151, 156, 158 S. domestica L . , 149, 156, 157 S. torminalis (L.) Crantz, 149, 151, 156 Sorghum (Gramineae), 211 S. arundinaceum (Desv.) Stapf, 108 S. purpureo-sericeum (Hochst. ex A . Rich.) Asch. & Schweinf., 108 Sorindeia (Anacardiaceae), 74 Soulamea terminalioides Baker (Simaroubaceae), 257 Soyauxia grandifolia Gilg & Stapf (Medusandraceae), 77 Spartium (Leguminosae: Papilionoideae), 147 S.junceum L . , 226 Spartocytisus nubigenus W e b b & Berth. (Leguminosae: Papilionoideae), 248 Spathionema (Leguminosae: Papilionoideae), 111 Spathodea campanulata P. Beauv. (Bignoniaceae), 182 Spergularia marítima (Hill) Druce (Caryophyllaceae), 266 Sphagnum (Sphagnaceae), 45, 84, 237, 266 Sphenopus (Gramineae), 266 S. divaricatus (Gouan) Reichenb., 228, 230 S. gouanii Trin.: see S. divaricatus Spirostachys africana Sond. (Euphorbiaceae), 95, 96, 200, 201 Spondianthus preussii Engl. (Euphorbiaceae), 83, 181 Spondias mombin L . (Anacardiaceae), 109 Sporobolus (Gramineae), 116,126 S. barbigerus Franchet: see S. subtilis S. centrifugus Nées, 239 S. discosporus Nées, 194 S. durits Brongn., 254 S. festivus Hochst. ex A . Rich., 128, 209, 210, 243 S. fimbriatus Nées, 194 S. humifusus (Kunth) Kunth, 212, 213
S. infirmus M e z , 84 S. ¡ociados (Trin.) Nées, 126 S. kentrophyllus (K. Schum.) W . D . Clayton, 126 S. nitens Stent, 267 S.pyramidalisP. Beaüv., 101, 191 5. robustus Kunth, 144, 223, 267 S. sanguineus Rendle, 84 S. spicatus (Vahl) Kunth, 116, 252, 267 S. subtilis Kunth, 101 S. subulatus Hackel ex Scott Elliot, 239 S. tenellus (Sprengel) Kunth, 267 S. testudinum Renvoize, 259 S. virginicus (L.) Kunth, 253, 259, 263, 267 Stachys spathulata Burchell ex Benth. (Labiatae), 194 Stadmannia oppositifolia Poir. (Sapindaceae), 257 Stangeria (Stangeriaceae), 199 5. eriopus (Kunze) Bâillon, 200 Stapelia (Asclepiadaceae), 139 Stapfiella (Turneraceae), 162 Stathmostelma pauciflorum (Klotzsch) K . Schum. (Asclepiadaceae), 100 S. welwitschii Britten & Rendle, 100 Statice (Plumbaginaceae), 266 S. cyrtostachya Boiss. & Reut.: see Limonium cymuliferum Staudtia stipitata W a r b . (Myristicaceae), 79, 85,172 Stauracanthus boivinii (Webb) S a m p . : see Ulex boivinii Steganotaenia araliacea Hochst (Umbelliferae), 96, 98, 99, 105 Stemonocoleus (Leguminosae: Caesalpinioideae), 74 Stenochlaena tenuifolia (Desv.) Moore (Blechnaceae), 200 Stenocline (Compositae), 238 Sterculia (Sterculiaceae), 187 S. africana (Lour.) Fiori, 114, 141 5. appendiculata K . Schum., 186, 188, 189 S. oblonga Masters, 78, 79 S. quinqueloba (Garcke) K . Schum., 173,174 S. rhinopetala K . Schum., 79 S. rhynchocarpa K . Schum., 114, 189 S. rogersii N . E . Br., 96 S. setigera Del., 62, 90,105, 107, 145, 211 S. stenocarpa H . Winkler, 114, 116 S. tragacantha Lindley, 74, 82, 178, 253 Stereospermum acuminatissimum K . Schum. (Bignoniaceae), 82 S. euphorioides D C , 241, 243 S. kunthianum C h a m . , 62, 83, 85, 105, 107, 173, 174, 189, 210,211,212 S. variabile H . Perrier, 243 Sticherus flagellaris (Bory) St John (Gleicheniaceae), 236 Stipa (Gramineae), 230 S. capensis Thunb., 222, 227, 228 S. parviflora Desf., 222 S. retorta Cav.: see S. capensis S. tenacissima L., 40, 51, 52, 149, 153, 155, 216, 226,229, 230 S. tortilis Desf.: see S. capensis Stipagrostis (Gramineae), 137, 140, 142, 144, 193 S. acutiflora (Trin. & Rupr.) de Winter, 212 S. amabilis (Schweick.) de Winter, 191, 193 S. brevifolia (Nées) de Winter, 141, 142 S. ciliata (Desf.) de Winter, 141, 142, 193, 204, 221 S. gonatostachys (Pilger) de Winter, 142 S. hermannii (Mez) de Winter, 142 S. hirtigluma (Steud. ex Trin. & Rupr.) de Winter, 113, 143, 144, 204 S. hochstetterana (Beck, ex Hackel) de Winter, 143, 145 S. namaquensis (Nées) de Winter, 141 S. namibensis de Winter, 142
Index of plant names
S. obtusa (Del.) Nees, 141,142,193, 221, 222 S. plumosa (L.) M u n r o ex T . Anderson, 221 S. pungens (Desf.) de Winter, 147, 204, 207, 216, 219, 220, 224, 230 S. ramulosa de Winter, 142 S. sabulicola (Pilger) de Winter, 142 S. subacaulis (Nees) de Winter, 143,144 S. uniplumis (Licht. ex R o e m . & Schult.) de Winter, 113, 141, 145, 193, 204 S. zitellii (Asch.) de Winter, 224 Stoebe (Compositae), 132,167, 238 S. passerinoides Willd., 258 Streptocarpus (Gesneriaceae), 165 Striga hermonthica (Del.) Benth. (Scrophulariaceae), 212 Strombosia grandifolia Hook.f. ex Benth. (Olacaceae), 85 S. scheffleri Engl., 164, 181, 187 S. ¡¡p., 253 Strychnos (Loganiaceae), 75 S. cocculoides Baker, 173 S. decussata (Pappe) Gilg, 200 S. henningsii Gilg, 90, 129, 174, 200 S. innocua Del.: see S. madagascariensis S. madagascariensis Poir., 85, 90, 99,105,189, 200, 211 S. mellodora S. Moore: see S. mitis S. mitis S. M o o r e , 122, 187 S. potatorum L.f., 90, 91, 98, 99, 182 S. pungens Solered., 85, 96, 97,173, 196 S. spinosa L a m . , 85,107, 189, 210 S. stuhlmannii Gilg: see S. potatorum S. usambarensis Gilg, 167 Stuhlmannia (Leguminosae: Caesalpinioideae), 186 Suaeda (Chenopodiaceae), 223, 224, 267 S. articulata Aellen, 267 S. fruticosa Forssk. ex J. F . Gmelin, 144, 223, 228, 230, 266 S. ifniensis Caball., 223, 229 S. mollis (Desf.) Del., 223, 229 5. monodiana Maire, 223 S. monoica Forssk. ex J. F . Gmelin, 31, 120, 224, 263, 267 S. plumosa Aellen, 144 S. vermiculata Forssk. ex J. F . Gmelin, 223 Suregada africana (Sond.) Kuntze (Euphorbiaceae), 200 S. procera (Prain) Croizat, 129, 167,181 S. zanzibarensis Bâillon, 188, 189 Sutera (Scrophulariaceae), 140 Swartzia madagascariensis Desv. (Leguminosae: Caesalpinioideae), 87, 97, 105, 173 Symphonia globulifera L.f. (Guttiferae), 83, 85, 181, 235, 236, 237, 253 Syzygium cordatum Hochst. ex Krauss (Myrtaceae), 91, 168, 181, 189,196,200,202,266 S. guiñéense (Willd.) D C , 85,181,189, 211 ss. afromontanum F . White, 90, 92, 164 ss. bamendae F . White, 253 ss. barotsense F . White, 91 ss. gerrardii (Harv. ex Hook.f.) F . White, 200 ss. guiñéense, 97, 99,105 ss. occidentale F . White, 82 S. owariense (P. Beauv.) Benth., 74, 91 S. sclerophyllum Brenan, 187 Tabernaemontana elegans Stapf (Apocynaceae), 189 T. johnstonii (Stapí) Pichón, 164, 167 T. stenosiphon Stapf, 253 Tagetes patula L . (Compositae), 252 Talbotiella (Leguminosae: Caesalpinioideae), 74
353
T. gentil Hutch. & Greenway, 176 Tamarindus indica L . (Leguminosae: Caesalpinioideae), 95,105, 107,117,129,182,188,189, 209, 241, 243, 251, 252, 255 Tamarix (Tamaricaceae), 47, 207, 216, 219 (and see footnote), 223, 224, 266, 267 T. aphylla (L.)Ka.isL, 113 T. 'articulata', 219, 220 T. canariensis Willd., 251 T. 'gallica', 219 ss. 'nilotica', 222 T. 'mannifera', 224 T. nilotica (Ehrenb.) Bunge, 113, 219 T. usneoides E . M e y . ex Bunge, 141, 144, 191 T. sp., 223 Tambourissa (Monimiaceae), 232, 235, 236, 258 T. gracilis Baker, 238 Tamus communis L . (Dioscoreaceae), 151 Tapiphyllum floribundum Bullock (Rubiaceae), 97 Tarchonanthus camphoratus L . (Compositae), 115, 135,191, 193, 195, 201 T. galpinii Hutch. & E . P . Phillips, 96 T. minor Less., 193 Tarenna graveolens (S. M o o r e ) Bremek. (Rubiaceae), 129,182 T. tuteóla (Stapf) Bremek., 90 T. neurophylla (S. M o o r e ) Bremek., 98, 99 Tarietia utilis (Sprague) Sprague (Sterculiaceae), 76 Taxus baccata L . (Taxaceae), 149,151, 152, 156, 247 Teclea (Rutaceae), 115, 167 T. gerrardii I. Verdoorn, 200 T. nobilis Del., 122, 129, 182, 211 T. simplicifolia (Engl.) I. Verdoorn, 115, 121,122,129 T. trichocarpa (Engl.) Engl., 129 Teline linifolia (L.) W e b b & Berth. (Leguminosae: Papilionoideae): see Genista linifolia T. monspessulana (L.) K . K o c h : see Cytisus monspessulanus T. stenopetala ( W e b b & Berth.) W e b b & Berth.: see Cytisus stenopetalus Teloschistes capensis (L.f.) M a l m e (Teloschistaceae), 142 Tephrosia (Leguminosae: Papilionoideae), 140, 220 T. gracilipes Guill. & Perr., 204 T. nubica (Boiss.) Baker, 204 T. obcordata (Lam. ex Poir.) Baker, 204 T. quartiniana Cuf., 204 T. uniflora Pers., 206 Terminalia (Combretaceae), 110, 111 T. avicennioides Guill. & Perr., 105, 107,108 T. bentzoe Pers., 257, 258 T. benzoin L.f.: see T. bentzoe T. boivinii Tul., 259 T. brachystemma W e l w . ex Hiern, 99 T. brownii Fresen., 107, 208, 209, 210, 211, 212, 213, 214 T. glaucescens Planch, ex Benth., 83, 85,105,107 T. laxiflora Engl., 85, 105,107, 108, 203, 209, 210, 212 T. macroptera Guill. & Perr., 105, 107, 108 J. mollis Lawson, 95, 129,173, 174 T. orbicularis Engl. & Diels, 114 T. párvula P a m p a n . , 114 T, prunioides Lawson, 95, 96, 143,144, 191 T. sambesiaca Engl. & Diels, 117,186, 188 T. sericea Burchell ex D C , 95, 96, 97,191,193,194, 201 T. seyrigii ( H . Perrier) Capuron, 243 T. spinosa Engl., 114, 188, 189 T. stuhlmannii Engl., 116 T. subserrata H . Perrier, 242 T. superba Engl. & Diels, 57, 75, 78, 79, 81
354
Index of plant names
Tessmannia (Leguminosae: Caesalpinioideae), 74 T. burttii H a r m s , 91 Tetraberlinia (Leguminosae: Caesalpinioideae), 74 T. bifoliolata (Harms) H a u m a n , 77 T. polyphylla (Harms) J. Léonard, 77 T. tubmaniana J. Léonard, 77 Tetracera (Dilleniaceae), 241 Tetraclinis (Cupressaceae), 147, 227 T. articulata (Vahl) Masters, 48, 146, 149, 150, 151, 152, 153,154,155,157,227,228,229 Tetragonia (Aizoaceae), 137, 193 T. reduplicata W e l w . ex Oliver, 145 Tetrapleura tetraptera (Schumach. & Thonn.) Taubert (Leguminosae: Mimosoideae), 74, 82,181,253 Tetrapogon cenchriformis (A. Rich.) W . D . Clayton (Gramineae),209,210,212 T. tenellus (Roxb.) Chiov., 145 Tetrapterocarpon (Leguminosae: Caesalpinioideae), 232 T. geayi Humbert, 242 Tetraria (Cyperaceae), 132 Tetrorchidium didymostemon (Bâillon) Pax & Hoffm. (Euphorbiaceae), 80 Teucrium (Labiatae), 147 T.fruticansL., 154, 158 T. polium L . , 152, 154 Thalia welwitschii Ridl. (Marantaceae), 108 Thamnochortus erectus (Thunb.) Masters (Restionaceae), 135 T. spicigerus (Thunb.) R . Br., 135 Thelypteris confluens (Thunb.) Morton (Thelypteridaceae), 266 T. striata (Schumach.) Schelpe, 265 Themeda quadrivalvis (L.) Kuntze (Gramineae), 243, 255 T. triandra Forssk., 32, 100, 116, 121, 126, 127, 128, 129, 137, 168,169,191, 193, 194,195, 196, 202, 208, 211 Thesium (Santalaceae), 238 Thespesia danis Oliver (Malvaceae), 188,189 Thonningia (Balanophoraceae), 75 T. sanguínea Vahl, 93 Thunbergia crispa Burkill (Acanthaceae), 99 T. guerkeana Lindau, 114 Thylachium africanum (Capparidaceae), 188 T. thomasii Gilg, 114 Thymelaea lythroides Barratte & M u r b . (Thymelaeaceae), 152 T. microphylla Coss. & Durieu, 230 T. nitida Desf., 230 Thymus caespititius Brot. (Labiatae), 247 Tibestina (Compositae), 218 Tieghemella (Sapotaceae), 74 T. africana Pierre, 77 T. heckelii Pierre ex A . Chev., 77 Timonius seychellensis Summerhayes (Rubiaceae), 257 Tina (Sapindaceae), 232 T. isoneura Radlk., 237 Tinnea aethiopica Kotschy & Peyr. (Labiatae), 115 Titanopsis (Aizoaceae), 140 Tournefortia argéntea L.f. (Boraginaceae), 257 Trachylobium verrucosum (Gaertner) Oliver: see Hymenaea verrucosa Trachypogon spicatus (L.f.) Kuntze (Gramineae), 100, 194, 196, 239 T. thollonii Stapf, 101 Traganopsis glomerata (Maire) Wilczek (Chenopodiaceae), 229 Traganum nudatum Del. (Chenopodiaceae), 223 Tragus berteronianus Schult. (Gramineae), 194, 243 T. koelerioides Asch., 194, 195 T. racemosus (L.) Ail., 194, 204, 206, 207
Trapa natans L . (Trapaceae), 265 Treculia africana Decne. (Moraceae), 91,172, 187, 253 Trema guineensis (Schumach. & Thonn.) Ficalho (Ulmaceae): see T. orientalis T. orientalis (L.) Bl., 25, 80, 83, 211 Trianthema hereroensis Schinz (Aizoaceae), 142 Tribulocarpus dimorphanthus (Pax) S. M o o r e (Aizoaceae), 113 Tribulus (Zygophyllaceae), 144 T. terrestris L . , 194, 206, 212 T. zeyheri Sond., 141,143 Tricalysia allenii (Stapf) Brenan (Rubiaceae), 90 Trichilia dregeana Sond. (Meliaceae), 187, 200 T. emética Vahl, 91, 95, 96,105, 117,188, 200, 201 T. grandifolia Oliver, 253 T. prieuriana A d r . Juss., 74, 90 Trichocalyx (Acanthaceae), 111 Trichocaulon (Asclepiadaceae), 139 T. clavatum (Willd.) H . Huber, 142 T. dinteri Berger: see T. clavatum T. pedicellatum Schinz, 142 Trichocladus (Hamamelidaceae), 162 T. ellipticus Eckl. & Zeyh., 167, 181 Tricholaena monachne (Trin.) Stapf & C . E . Hubbard (Gramineae), 145 Trichomanes tnannii H o o k . (Hymenophyllaceae), 82 Trichoneura grandiglumis (Nées) E k m a n (Gramineae), 194 Trichoscypha (Anacardiaceae), 74 Trifolium (Leguminosae: Papilionoideae), 147,157,166 Triglochin palustris L . (Juncaginaceae), 155 Trilepisium madagascariense D C . (Moraceae), 79, 172, 187, 188,253 Trimeris (Campanulaceae), 254 T. scaeviolifolia (Roxb.) Mabberley, 254 Triplocephalum holstii O . Hoffm., (Compositae), 267 Triplochiton scleroxylon K . Schum. (Sterculiaceae), 57, 79, 81 Tripogon leptophyllus (A. Rich.) Cuf. (Gramineae), 211 T. minimus (A. Rich.) Hochst. ex Steud., 210 Triraphis andropogonoides (Steud.) E . P. Phillips (Gramineae), 194 Tristachya eylesii Stent & Rattray (Gramineae): see T. nodiglumis T. hispida (L.f.) K . Schum.: see T. leucothrix T. leucothrix Nées, 194, 196, 202 T. nodiglumis K . Schum., 173 Tristemma incompletum R . Br. (Melastomataceae), 266 Triumfetta annua L . (Tiliaceae), 90 Trochetia erythroxylon (G. Forster) Benth. (Sterculiaceae), 254 T. melanoxylon (Aiton) Benth. & Hook.f., 254 Turraea floribunda Hochst. (Meliaceae), 200 T. ghanensis J. B . Hall, 176 T. glomeruliflora H a r m s , 253 T. holstii Giirke, 181 T. mombassana Hiern ex C . D C , 115, 121 T. nilotica Kotschy & Peyr., 182 T. obtusifolia Hochst., 200 T. oppositifolia (Cav.) H a r m s , 258 Turraeanthus (Meliaceae), 74 Typha (Typhaceae), 55,219 T. australis Schumach. & Thonn., 191,222,265 T. latifolia L., 21», 265 Typhonodorum lindleyanum Schott (Araceae), 188, 236 Uapaca (Euphorbiaceae), 99,106 U. bojeri Bâillon, 237 U. chevalieri Beille, 82
Index of plant names
U. guineensis Muell.-Arg., 83, 85, 91, 181 U. heudelotii Bâillon, 83 U. kirkiana Muell.-Arg., 99 U. nítida Muell.-Arg., 173,189 U. pilosa Hutch., 99 U. sansibarica Pax, 173, 189 U. togoensis Pax, 85, 105 Ulex boivinii W e b b (Leguminosae: Papilionoideae), 151,152 U. europaeus L., 254 Ulmus campes tris L . (Ulmaceae), 149, 150 Umtiza (Leguminosae: Caesalpinioideae), 199 U. listerana Sim, 200 Uncarina (Pedaliaceae), 242 Urelytrum giganteum Pilger (Gramineae), 174 U. squarrosum Hackel, 239 Urena lobata L. (Malvaceae), 206 Urginea (Liliaceae), 158 U. marítima (L.) Baker, 152,160 Ursinia (Compositae), 141 Usnea (Parmeliaceae), 99,142,181, 237 Utricularia (Lentibulariaceae), 108, 265 U. foliosa L., 265 U. gibba L., 266 Uvaria chamae P. Beauv. (Annonaceae), 81,178 U. leptocladon Oliver, 189 Uvariodendron anisatum Verde. (Annonaceae), 167 Vaccinium (Ericaceae), 168,236,237,238 V. cylindraceum Smith, 247 Vahlia geminiflora (Del.) Bridson (Vahliaceae), 204 Vallisneria aethiopica Fenzl (Hydrocharitaceae), 265 V. spiralis L . , 265 Vangueria venosa Hochst. ex Del. (Rubiaceae), 207 Vangueriopsis lanciflora (Hiern) Robyns (Rubiaceae), 97, 99 Vanilla roscheri Reichenb.f. (Orchidaceae), 114 Valeria (Dipterocarpaceae), 257 V. seychellarum Dyer, 257 Vella mairei Humbert (Cruciferae), 158 Venidium (Compositae), 142 Vepris heterophylla (Engl.) Letouzey (Rutaceae), 178 V. undulata (Thunb.) I. Verdoorn & C . A . Smith, 200 Verbascum capitis-viridis Huber-Mor. (Scrophulariaceae), 252 Vernonia (Compositae), 236, 237, 238 V. amygdalina Del., 253 V. auriculifera Hiern, 130 V. brachycalyx O . Hoffm., 182 V. confería Benth., 80 V. oligocephala ( D C . ) Schultz.-Bip. ex Walp., 194 Verschaffeltia (Palmaceae), 257 V. splendida H . A . Wendl., 257 Vetiveria fulvibarbis (Trin.) Stapf (Gramineae), 178 V. nigritana (Benth.) Stapf, 108 Viburnum lantana L. (Caprifoliaceae), 151 V.tinusL., 151,157,247 Vicia (Leguminosae: Papilionoideae), 147,157 Vigna luteola (Jacq.) Benth. (Leguminosae: Papilionoideae), 265 V. unguiculata (L.) Walp., 252 Viola (Violaceae), 236 V. arborescens L., 154 V. palustris L., 155 Viridivia (Passifloraceae), 87 Visnea (Theaceae), 246 V. mocanera L.f., 246, 247 Vitellariopsis marginata (N. E . Br.) Aubrév. (Sapotaceae), 200 Vitex (Verbenaceae), 237
355
V. agnus-castus L., 147, 219 V. doniana Sweet, 83, 85,105, 107 V. humbertii Moldenke, 238 V. madiensis Oliver, 85 V. mombassae Vatke, 189 Vitis (Vitaceae), 159 Vinaria elongata Swartz (Vittariaceae), 235 Voacanga thouarsii R o e m . & Schult. (Apocynaceae), 83, 181, 188,200 Volkensinia (Amaranthaceae), 111 Vossia cuspidata Griff. (Gramineae), 55, 100, 265, 266 Vulpia bromoides (L.) S. F . Gray (Gramineae), 211 Wahlenbergia (Campanulaceae), 140, 254 W. angustifolia (Roxb.) A . D C , 254 W. linifolia (Roxb.) A . D C , 254 Walafrida (Scrophulariaceae), 135, 140, 194 W. densiflora Rolfe, 194 W.saxatilis Rolîe, 194,195 Waltheria indica L . (Solanaceae), 206 Warburgia saluions (Bertol.f.) Chiov. (Canellaceae), 167,181 W. ugandensis Sprague: see W. salutaris Warionia (Compositae), 218 Weinmannia (Cunoniaceae), 236, 237, 238, 255 Wellstedia (Boraginaceae), 113 Welwitschia bainesii (Hook.f.) Carrière (Welwitschiaceae), 53, 94,136, 137, 143,144,145, 191 W. mirabilis Hook.f.: see W. bainesii Wiborgia sericea Thunb. (Leguminosae: Papilionoideae), 134 Widdringtonia (Cupressaceae), 135, 167 W. cedarbergensis J. A . Marsh, 134 W. cupressoides (L.) Endl., 51, 134,161,165, 166 W. nodiflora (L.) Powrie: see W. cupressoides W. schwarzii (Marloth) Masters, 134 W. whytei Rendle: see W. cupressoides Willdenowia striata Thunb. (Restionaceae), 135 Wissmannia (Palmaceae), 111 W. carinensis (Chiov.) Burret, 113 Withania frutescens (L.) Pauquy (Solanaceae), 152, 227, 228 Wolffia arrhiza (L.) Horkel ex W i m m . (Lemnaceae), 265 Xanthocercis (Leguminosae: Papilionoideae), 87 X. zambesiaca (Baker) Dumaz-le Grand, 95 Xerocladia (Leguminosae: Mimosoideae), 137 Xeroderris stuhlmannii (Taubert) Mendonça & E . P. Sousa (Leguminosae: Papilionoideae), 95,106 Xeromphis nilotica (Stapf) Keay (Rubiaceae), 188 X. rudis (E. Meyer ex Harv.) Codd, 201 Xerophyta (Velloziaceae), 238, 239, 242 X. dasylirioides Baker, 238 X. humilis (Baker) T . Durand & Schinz, 113 Xerosicyos (Cucurbitaceae), 232, 242 Ximenia (Olacaceae), 98 X. americana L., 91, 95,107, 253 X. caffra Sond., 95 Xylia hildebrandtii Bâillon (Leguminosae: Mimosoideae), 241 Xylocalyx (Scrophulariaceae), 111 Xylocarpus (Meliaceae), 261 X. granatum Koen., 259, 261, 263, 264 X. moluccensis (Lam.) M . J . R o e m . , 259, 261 Xylopia aethiopica (Dunal) A . Rich. (Annonaceae), 74, 81, 91, 253 X. holtzii Engl.: see X. parviflora X. odoratissima W e l w . ex Oliver, 97 X. parviflora (A. Rich.) Benth., 188
356
Index ofplant names
Xylopia aethiopica (Dunal) A . Rich. (Annonaceae)—contd X. rubescens Oliv., 91 Xymalos (Monimiaceae), 162 X. monospora (Harv.) Bâillon, 122, 164, 165,181, 187, 199 Xyris (Xyridaceae), 84, 108 Zanha africana (Radlk.) Exell (Sapindaceae), 97 Z . golungensis Hiern, 109 Zanthoxylum capense (Thunb.) Harv. (Rutaceae), 200, 201 Z . chalybeum Engl., 189 Z . davyi (I. Verdoorn) Waterman, 199 Z . gilletii (de W ü d . ) Waterman, 81, 253 Z . trijugum (Dunkley) Waterman, 90 Z . xanthoxyloides (Lam.) ?, 109,178 Zea mays L . (Gramineae), 252 Zenkerella capparidacea (Taubert) J. Léonard (Leguminosae: Caesalpinioideae), 187 Z/7fa(Cruciferae),218 Z . spinosa (L.) Prantl, 221, 222, 224 Ziziphus (Rhamnaceae), 98, 208, 210 Z . abyssinica Hochst. ex A . Rich., 95, 106, 210
Z . lotus (L.) Desf., 147, 149, 158, 160, 204, 219, 225, 226, 227,228,231 Z . mauritiana L a m . , 105, 211, 219, 252 Z . mucronata Willd., 95, 96, 105, 106, 107, 129, 137, 141, 189,193,194,195,200,201 Z . pubescens Oliver, 129 Z . spina-christi (L.) Desf., 209, 210, 211, 212, 214 Z . zeyherana Sond., 194 Zornia glochidiata Reichb. ex D C . (Leguminosae: Papilionoideae), 212 Zygophyllum (Zygophyllaceae), 137, 140, 223 Z . album L . , 223, 224, 230 Z . cornutum Coss., 223 Z . gaetulum Emberger & Maire, 223, 229 Z . hildebrandtii Engl., 116 Z . morgsana L . , 135 Z . orbiculatum W e l w . ex Oliver, 144, 145 Z . retrofractum Thunb., 142 Z . simplex L . , 143,144, 204, 252 Z . stapfii Schinz, 142, 144 Z . waterlotii Maire, 223
ERRATA*
p. 24, explanation of Fig. 3, line 16: replace area by period p. 40L, line 21 : replace 1943 by 1934 p. 42, Table 2, column 4,line 15: replace XVII by XVIII p. 46, Table 3, column 4,line 21: replace \3by 14 p. 61, Mapping unit 28, column 1, line 2: replace 26 ¿y 28 p. 66, Mapping unit 76, column 2, line 7: replace XVII 6y XVIII p. 94L, line 9: replace Drumond ¿»y D r u m m o n d p. 104, Fig. 7: replace 5. Imatong Mts.ôy 5. Didinga Hills p. 107L, line 20: replace tomentosa by angolensis p. 122L, line 11 : replace most by more p. 122R,line 43: replace places by phases p. 135R, line 2: replace May tenus by *Maytenus p. 135R, line 3: replace Chionanthus by *Chionanthus p. 135R, lines 18-19: delete, Chionanthus (Linociera)foveolatus,May tenus heterophylla p. 141L, line 11 : replace besóme by become p. 1S8R, line 51 : replace Rhamus by Rhamnus p. l72R,line 24: replace west by east;replace north-west by north-east p. 172R, line 31 : replace west by east p. 192, top left-hand corner of map: replace Roman I by Arabic 1 p. 194R, line 47: replace karoo by karroo p. 199R, line 52: replace bachmanii by bachmannii p. 201L, line 42: replace above by below p. 205, caption to Fig. 21 : insert regional before transition p. 213L, line 11 : replace 000 by 23 p. 213L, line 12: replace 000 by 27 p. 213L, line 53: replace Erogrostis by Eragrostis p. 227L, line 40 and elsewhere: replace Drar by Dra p. 229R, line 54: replace europea by europaea p. 230R, line 49: delete comma afterferum p. 245, Fig. 24: replace Bioka by Bioko p. 249L, line 34: delete latitudes p. 250, Fig. 26: add 3 9 5 m to the island of Sta Luzia p. 2 51R, line 50 : replace Teixera by Teixeira p. 252L, line 2: replace bateta by batata p. 254R, line 12: replace potulacifolia by portulacifolia p. 259R, line 31 : replace molucensis by moluccensis p . 2 6 6 R , last line: replace Hannington by Bogoria p. 285L: after D E W I N T E R , insert B . p. 289R, line 23: replace herbeuses by herbeux p. 292L, line 43: replace stratioides by stratiotes p. 294R, line 5: before La insert 1927ft p. 296R, line 41 : replace V A N C O S T E N by V A N O O S T E N p. 300R, line 36: replace chorologique by chorologiques p. 301R, line 49: replace Amadoua by Adamaoua p. 305L, line 12: replace Egypt, by Egypt. p. 307R, line 50: replace Brandenberg by Brandberg p. 314L, line 44: transpose Arlon, Belgium and Univ. Luxemb. p. 336L, line 61 : replace colora by colona p. 353R, line 26: replace Tarietia by Tarrietia p. 356L, line 13: replace question mark by Zepernick & Timler * L and R - left- and right-hand columns respectively.
The Vegetation of Africa, Paris, Unesco, 1983
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