Carcasson's African Butterflies: An Annotated Catalogue of the Papilionoidea and Hesperioidea of the Afrotropical Region

By PR Ackery, CR Smith and R Vane-Wright

The result of more than 20 years of research and collaboration by international butterfly experts, this book is the first comprehensive catalogue to the butterfly fauna of any major tropical region and, as such, provides a basic research tool for any worker with an interest in African butterflies. Covering 3593 recognised species in 300 genera, it deals with about 20% of the world butterfly fauna. Included are entries for all genus-group, species-group and infra-subspecific names applicable to the Afrotropical butterflies, a total of about 14 000 names. This work has a more wide-ranging appeal than a narrow taxonomic list, a volume that will be of value not only to taxonomists but to all biologists with an interest in Africa and its butterfly fauna.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Jan 1, 1995
• ISBN: 9780643102453

Book preview

CARCASSON’S

AFRICAN

BUTTERFLIES

An Annotated Catalogue of the Papilionoidea and Hesperioidea of the Afrotropical Region

Editors

P.R. Ackery, C.R. Smith & R.I. Vane-Wright

The Natural History Museum, London, U.K.

CSIRO Australia Cataloguing-in-Publication

Carcasson’s African Butterflies: an annotated catalogue of the Papilionoidea and Hesperioidea of the Afrotropical region editors, P.R. Ackery, C.R. Smith & R.I. Vane-Wright.

Includes bibliographic references and index.

ISBN 0 643 05561 4

1. Papilionoidea — Africa, East 2. Papilionoidea — Catalogs and collections 3. Hesperioidea — Africa, East 4. Hesperioidea — Catalogs and collections 1. Ackery, Phillip Roland. II. Smith, C.R. (Campbell R.) III. Vane-Wright, Richard Irwin. IV. Carcasson, R.H. (Robert Herbert) V. CSIRO. VI. Natural History Museum (London) VII. Title: African Butterflies

595.789

© CSIRO Australia 1995

The intellectual content of this book, Carcasson’s African Butterflies: an annotated catalogue of the Papilionoidea and Hesperioidea of the Afrotropical region, remains the property of The Natural History Museum, London U.K.

This book is available from:

CSIRO Publications

PO Box 89 (314 Albert Street)

East Melbourne, Victoria 3002

Australia

Tel. [phone]      Int: +(613) 9418 7217

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email: [e-mail]

Managing Editor: Kevin Jeans

Cover and Layout Design: Anita Adams

Typesetter: Trevor Clarke

Production Manager: Jim Quinlan

DEDICATION

During the last 20 years, a number of outstanding specialists on African butterflies have died, including Bob Carcasson himself. We think particularly of Charlie Dickson, David Gifford and Dimitri Sevastopulo who were well known to us and who, together with Bob, provided inspiration to complete this work.

R.H. CARCASSON 1918-1982

For Bob Carcasson, the publication of his catalogue would have been the culmination of a lifetime’s interest in natural history, and Lepidoptera in particular. In 1971, he first approached Dick Vane-Wright with an incomplete draft manuscript, suggesting that it was a project on which we could fruitfully collaborate. By the end of 1974, Dick and Clive Huggins had completed the Papilionidae as a pilot project, and since then work has continued on re-drafting and up-dating Bob’s typescript. It is a matter of deep regret to us that Bob did not live to see the completion of his monumental work, but we are grateful for the continued interest and forbearance of his family (particularly George V. Carcasson) during what must surely be one of the longest gestation periods for any lepidopterological volume.

Bob Carcasson was born in Cheltenham, England, in 1918. Following the death of his father, Bob and his Italian mother moved to Florence, and it was in Italy that he qualified for an agricultural diploma. Here, too, began a friendship and collaboration with Roger Verity, author of the classic work on Italian butterflies. Bob arrived in Zimbabwe (then Southern Rhodesia) to farm, before joining the Forces and seeing service in Ethiopia, Somalia, North Africa and Italy. Demobilisation and marriage followed, and with his wife, Bee, he returned to Zimbabwe to farm in the Manicaland mountains.

Ten years of farming ended when he joined a pest control firm and was transferred to Kenya. In 1955, he succeeded E.C.G. Pinhey as entomologist at the Coryndon Museum (now the National Museum), and on Louis Leakey’s retirement, became Director of the Museum in 1961. Shortly after the award of his PhD by the University of Nairobi for work on the Sphingidae, Bob took up a new appointment as Chief Curator of the Centennial Museum in Vancouver. Leaving this post in 1971, he then travelled extensively in the Far East, working on a field guide to Indo-Pacific coral fishes. From 1973 until his retirement because of ill-health in 1979, he was Curator of Entomology at the Provincial Museum of British Colombia. Following the removal of one lung, due to cancer, Bob still found the time and motivation to publish his now standard Hand Guide to the Butterflies of Africa. Titles of his other major studies on African butterflies appear in the bibliography of this present volume. Only someone with his exceptional knowledge could have undertaken to survey the biogeography of the butterflies of the whole of Africa (1964, East African Wildlife Journal 2: 122-157). An edited version of this wide-ranging and pioneering work is reprinted in the introductory section (commences on page 17).

For us, this publication brings to an end some 20 years of labour. We hope that it is a fitting memorial to a charming, multi-talented man, while regretting that in its final version it did not have the benefit of his fine touch, personal knowledge and judgement.

Phillip Ackery

Campbell Smith

Dick Vane-Wright

CONTENTS

LIST OF CONTRIBUTORS

ACKNOWLEDGEMENTS

FOREWORD

INTRODUCTION

BIOLOGY

GENERAL BEHAVIOUR

EARLY STAGE BIOLOGY

GAZETTEER AND LIST OF REGIONAL WORKS

BIOGEOGRAPHY

BUTTERFLIES AND THE AFROTROPICAL REGION

AFRICAN REGIONAL GEOGRAPHY

AFRICAN CLIMATE AND VEGETATION

CLIMATIC CHANGES DURING THE LATE TERTIARY AND QUATERNARY

SPECIATION, SUBSPECIATION AND VICARIANCE

SEGREGATION MECHANISMS

RE-INTEGRATION OF DISJUNCT POPULATIONS

AFROTROPICAL FAUNISTIC REGIONS

Sylvan sub-region

Sub-region of open formations

Cape sub-region

Malgassy sub-region

Special habitats

Eclectic species

FAUNAL AFFINITIES

ANOMALOUS DISCONTINUITIES

DIVERSITY, ENDEMISM AND CONSERVATION

THE CATALOGUE

SCOPE

CONVENTIONS

NOMENCLATORIAL CHANGES

ABBREVIATIONS

HESPERIOIDEA

Hesperiidae

Coeliadinae

Pyrginae

Hesperiinae

PAPILIONOIDEA

Papilionidae

Papilioninae

Troidini

Papilionini

Lampropterini

Pieridae

Pseudopontiinae

Coliadinae

Pierinae

Euchloini

Pierini

Nymphalidae

Acraeinae

Danainae

Danaini

Euploeini

Satyrinae

Melanitini

Elymniini

Satyrini

Argynninae

Melitaeinae

Nymphalinae

Limenitinae

Biblini

Cyrestini

Neptini

Limenitini

Euthalini

Charaxinae

Apaturinae

Libytheinae

Lycaenidae

Lipteninae

Pentilini

Liptenini

Liphyrinae

Miletinae

Miletini

Spalgini

Lachnocnemini

Theclinae

Amblypodini

Oxylidini

Cheritrini

Aphnaeini

Iolaini

Hypolycaenini

Deudorigini

Lycaeninae

Polyommatinae

Lycaenesthini

Polyommatini

Riodinidae

Hamearinae

ADDENDUM

BIBLIOGRAPHY

INDEX

LIST OF CONTRIBUTORS

PHILLIP R. ACKERY

Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

RAMNIK ARORA

formerly Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

ROBERT H. CARCASSON (the late)

British Columbia Provincial Museum, Victoria, British Columbia, V8W 1A1, Canada

CINDY M. COGAN (formerly North)

formerly Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

CLIVE F. HUGGINS

formerly Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

ROBERT L. SMILES

formerly Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

CAMPBELL R. SMITH

Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

HELEN TAYLOR

formerly Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

RICHARD I. VANE-WRIGHT

Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

ACKNOWLEDGEMENTS

For some 20 years the staff associated with the butterfly collections at The Natural History Museum (BMNH) have been contributing to this work. The editors are particularly grateful to our former colleagues, Ramnik Arora, Cindy Cogan, Clive Huggins, Richard Sandifer, David Shuker, Bob Smiles and Helen Taylor. Bob Smiles was closely associated with this project for 10 years, not only contributing substantially to the text but also nursing his colleagues through the (for us) difficult transition between stone-age typescript and comparatively modern technology. Other authorities on African butterflies have given generously of their expertise – we owe a debt of thanks to Jonathan Ball, Georges Bernardi, Bernard D’Abrera, Rienk de Jong, the late Charlie Dickson, Jacques Hecq, Jan Kielland, Torben Larsen, David Lees, Jacques Pierre, Arthur Rydon and Gerry Tremewan. Many others have helped with advice on particular problems, including S.C. Collins, the late C.F. Cowan, A.J. Gardiner, D.L. Hancock, A. Heath, S.F. Henning, G.A. Henning, K. Fiedler, D.P. Knoop, E.L. Pringle, B. Turlin and M.B. Usher. Many colleagues in London assisted us over the years, notably long-suffering librarians and members of our photographic unit. The former Keeper of Entomology at The Natural History Museum, Prof. Laurence Mound, and his predecessor, Dr Paul Freeman, must at times have despaired of ever seeing any return on all this staff time. However, they kept these thoughts to themselves (mostly!) and we are grateful for their forthright backing of the project. Moral support has also come from George Carcasson and, more recently, the current Keeper of Entomology, Dr Richard Lane, and Prof. Bob May.

When this work was first started, publication of the results would have been little more than a formality. Twenty years later, in a world said to be crying out for comprehensive accounts of animals and plants, publication of such a book is anything but guaranteed. We therefore gladly acknowledge the Natural Environment Research Council for a grant which subvented some of the design costs, and offer our most grateful thanks to CSIRO for undertaking the enormous task of design, publication and distribution. It is a pleasure to record, in particular, the enthusiasm, patience and expertise of Trevor Clarke, Kevin Jeans and Anita Adams. Finally, we must thank Ebbe Nielsen (Australian National Insect Collection, Canberra) for faith in us, the project, the product and his belief that publication of this volume was not only possible, but also worthwhile.

FOREWORD

The distribution of taxonomic effort among taxonomic groups — the taxonomy of taxonomists – is very peculiar, reflecting past fashions more than present needs. Even among named and recorded species, the ratio of numbers of taxonomists to numbers of species is 100 times greater for vertebrate species than for invertebrate species, and 10 times greater for species of vascular plants than for invertebrates. These disparities are even more extreme when you recognise that most vertebrate species have indeed been recorded, along with an estimated 90% of the world’s vascular plants, whereas credible estimates suggest that the true total of insect and other invertebrate species is 3 to 30 times the number so far recorded. How are we to optimise our efforts to conserve the biological diversity we inherited, if we lack synoptic catalogues of what we have today? How are we to understand the causes and consequences of biological diversity – the relation between diversity and ecosystem structure and function — if we do not even have a list of what species are where?

To my mind, one of the larger mysteries of the human psyche is that we have long been fascinated by the physics of the very large and the very small – the fate of the universe and the secrets of the atom but only recently have begun systematically to study the other living things with which we share our planet. The astronomers have long had computerised catalogues of stars, galaxies, and other objects. All manner of fundamental understanding — grand patterns in the structure of the universe — is currently emerging from computer studies of these catalogues, which are only possible if such a database exists. And yet we still lack coordinated, synoptic catalogues of those species which have been named and recorded. We cannot even say how many species have been named and recorded. This simple, factual number is implicit in the sum of all existing catalogues (some now computerised, some even available as CD-ROM, but others still on aged file cards); but, lacking coordination, we lack the number. This is not a question of how many stamps there are: as the astronomers’ studies show, yesterday’s databases are the raw stuff for today’s recognition of patterns, leading to tomorrow’s fundamental understanding. Which brings me to Carcasson’s African Butterflies. It is a splendid achievement. Not just names, but information about geographical distribution, habitats, larval host-plants, and other ecological correlates of some 3600 species in 300 genera (and 14 000 names, which highlights the larger issues of synonymy, which further complicate things).

The book is much more than a triumph of scholarship in taxonomy and systematics. As human numbers, with all their associated environmental impacts, continue to grow, compilations of information like this one will assume ever-greater importance. Butterflies are often indicator species, whose declining populations give early warning of the effects of environmental changes on particular habitats and ecosystems, before such effects become more widely noticeable.

Butterflies are, of course, privileged or elite among the insects. They have the status, almost, of honorary birds (the Royal Society for the Protection of Birds in the UK has roughly one million members; the Butterfly Conservation Society has around ten thousand; no other group of insects draws on anything like this support). On the one hand, this gives Carcasson’s African Butterflies special importance; almost one fifth of the world’s butterflies are to be found in it. On the other hand, I believe that we need, to paraphrase a slogan of the 1960s, "two, three, many Carcasson’s African Butterflies". We need such synoptic catalogues, replete with ecological information about distributions, habitat, and so on, not just for the furries, featheries, and their cousins-in-sentiment the butterflies, but for the unnumbered hosts of less romantic beedes and other insects and invertebrates. Ultimately, we could discover too late that the continued functioning of the thin skin of life on the surface of our planet depends on these small things.

Robert M May

Oxford University and Imperial College, London

INTRODUCTION

The close of the nineteenth century saw the publication of the first monograph of the Afrotropical butterflies, P.O.C. Aurivillius’ monumental Rhopalocera Aethiopica (1898-9). This volume, still relevant today, was the forerunner of the few key works devoted to the systematics of the entire Afrotropical butterfly fauna. Such was Aurivillius’ expertise, that he contributed virtually all the text for the African volume of Die Gross-Schmetterlinge der Erde (Seitz, A. Ed., 1908-25). Although now much out-dated, this work provides a standard that has never been superseded. However, the checklists of Peters (1952) and Carcasson (1981), used in conjunction with D’Abrera’s monograph (1980), incorporate more recent developments and provide a generalized, but readily available, overview of the Afrotropical butterfly fauna.

This present work is the first comprehensive catalogue to the butterfly fauna of any major tropical region. Covering some 3600 recognised species and 300 genera, it deals with approximately 20% of the World butterfly fauna. Included are entries for all genus-group, species-group and infra-subspecific names applicable to the Afrotropical butterflies, a total of about 14 000 names. Within the catalogue itself, details will be found of distribution and larval hostplants, but under ‘Biology’ below we provide more general reviews of bionomics, early stage biology, the faunistic literature and biogeography. Thus we have endeavoured to provide a work of more wide-ranging appeal than a narrow taxonomic list, a volume that will be of value not only to taxonomists but to all biologists with an interest in Africa and its butterfly fauna.

BIOLOGY

The Afrotropical Region does not boast a 19th century giant, a Bates or a Wallace, as the local father of butterfly biology (although Roland Trimen, 1840-1916, could perhaps lay claim to that honour). Indeed, in some respects the African butterfly fauna continues to be neglected, in comparison to the Neotropics at least. While progress in the fertile fields of population and community biology has forged ahead in recent decades, few developments have their origins in Africa. But in other areas, African butterflies have been at the forefront, both historically and to the present day. Such topics as mimicry, migration, polymorphism, polyphenism, courtship and communication have all been profitably amplified through studies of African species and faunas. And with a significant wealth of national faunistic studies and local faunal lists, no comparable region is better served by the available literature. Below, we highlight the fields of general behaviour, early stage biology and biogeography, and the faunistic literature.

GENERAL BEHAVIOUR

The available faunistic studies carry a wealth of anecdotal information relating to habits, habitat preferences, range and local abundance. This is supplemented by a surprisingly small number of more specialized works. For anyone intertested in general bionomics, the classic accounts by Longstaff (1909, 1912), together with the all-embracing Seitz (1908-25), surely provide ideal starting points, in turn supplemented by D.F. Owen’s Tropical Butterflies (1971). Other less well-known works might also be consulted profitably: Birket-Smith (1960), Owen & Owen (1972, 1973), Wiley & Hudson (1943), Larsen et al. (1981) and Jackson (1961), and for the Lycaenidae alone, Lamborn (1914), Farquharson (1922) and Clark & Dickson (1971). Such comparative paucity reflects the state of our knowledge of African butterflies in the field, the extent to which collecting in transient expeditions has taken precedence over recorded observations of the butterflies themselves.

EARLY STAGE BIOLOGY

The primary role of butterfly larval foodplants in the founding studies of co-evolution (Ehrlich & Raven, 1965) and ecological biochemistry (Brower & Brower, 1964) has focussed attention on the wealth of available host data relating to butterflies (Ackery, 1988). We have throughout the catalogue noted the larval hostplants, in abbreviated form (see also Ackery, 1991), when they have come to our attention. This information has been drawn from a manuscript list (Ackery & Rajan, unpublished) in The Natural History (BMNH) Library, itself a compilation from various published sources. Those seeking more precise data should particularly consult Seitz (1908-25), which includes much valuable historical information, Platt (1921) for the South African fauna, van Someren (1974b) and Sevastopulo (1975) for eastern Africa, and Vinson (1938) for the Mascarene Islands. In addition, many of the national faunal monographs, especially Pinhey (1949b; 1965), Swanepoel (1953a), Gifford (1965), Pennington (1978), Migdoll (1988), Kielland (1990b) and Larsen (1991) contain copious details, again relating primarily to southern and eastern Africa. For western areas, comparable data are less readily available, usually being scattered in various journals, often in the form of annotated faunal lists (e.g. Birket-Smith, 1960, for Cameroon; Owen & Owen, 1972, 1973, for Sierra Leone) or more detailed studies of butterfly life cycles (see below).

The plant families most favoured as larval hosts by African butterflies include the Fabaceae, Poaceae, Euphorbiaceae, Sapindaceae, Lamiaceae and Acanthaceae (Ackery, 1991), a pattern of preference reflected in Australia (Symon, 1980) and probably elsewhere in the tropics. Members of endemic African families, or families of apparent ‘southern’ affinity, suffer relatively little predation, the Capys/Proteaceae and Poecilmitis/Zygophyllaceae associations being the two most outstanding exceptions. Analyzing the 41 families and 282 genera that comprise the Capensis floristic element of southern Africa, Cottrell (1978) found that members of only 12 genera, representing nine families, have been recorded as butterfly hosts.

More unusual larval resources are restricted to the lycaenid butterflies (Hinton, 1951; Eliot, 1973; Cottrell, 1984; Henning, 1983b). Species of Lipteninae (Alaena, Baliochila, Cooksonia, Durbaniopsis, Hewitsonia, Mimacraea, Ornipholidotos, Pentila, Teriomima and Durbania) favour lichens and occasionally simple algae. Elsewhere in the family, aphytophagy is widespread, most particularly in the Liphyrinae and Miletinae, but also in some polyommatine and thecline species. Such larval resources are varied, including either homopterans or their secrections (Aslauga, Megalopalpus, Spalgis, Lachnoncnema, Thestor, Triclema — see Ackery, 1990), or ant-brood or ant-reguritations (Lepidochryspos, Spindasis, Euliphyra).

Considering that the Afrotropical butterfly fauna comprises over 3600 species, it is surprising that so few have pest status; best known, perhaps, are the Rutaceae-feeding swallowtails, many of which now include Citrus within their range of accepted hosts. Potentially, indigenous satyrines and lycaenids could also assume importance. Monocultures of cereals present a possible new larval resource for the Satyrinae although, apart from occasional records of Melanitis leda on rice, this threat apparently has not been realized. Similarly, the legume-feeding lycaenids as yet do not appear to present a serious problem but, with their caterpillars feeding within the seed pods, they could significantly affect agricultural yields. Elsewhere, and particularly well-documented, are the preferences of Acraea acerata for sweet-potato and Deudorix (Virachola) livia for pomegranate. Acraea neobule defoliates the leafy vegetable Corchorus olitorius, used in soups in Nigeria where it is considered one of the cheapest sources of mineral elements and vitamins (Ewete, 1990, as terpsichore). Owen (1971) notes further scattered records for other butterflies as pests, notably Appias epaphia (on brassicas), Colias electo (Lucerne and Castor Oil), Catopsilia florella (ornamental cassias and Senna), Acraea neobule (Hibiscus, Jute) and Deudorix lorisona (Coffee).

Although there is an ever-increasing awareness of the importance of early stages in butterfly classification, the paucity of preserved voucher specimens of eggs, larvae, pupae, and even larval hostplants, is a striking feature of all major museum collections. The interested worker is largely reliant on published descriptions, ranging from the most cryptic through to the incomparable illustrations of the late Gowan C. Clark. Such studies are frequently ‘faunistic’ rather than ‘systematic’ in nature, with particularly important contributions based on the West African fauna. The pioneering works of Haig in Nigeria (Haig, 1933, 1936a, b, 1937, 1938a, b- see also Rosevear, 1978), complemented by Seth-Smith’s (1938) work on Ghanaian butterflies, stood virtually alone for several decades. Only in comparatively recent times have similar studies become available for Ivory Coast (Blandin et al., 1975; Vuattoux & Blandin, 1977, 1979; Vuattoux, 1988; Pierre & Vuattoux, 1978; Pierre-Baltus, 1978) and Zaire, Rwanda and Burundi (Fontaine, 1981a, b, 1982-3, 1985b, 1986, 1988a-e).

Gowan Clark’s illustrations illuminate the standard works on the early stage biology of South African butterflies (van Son, 1949a, 1955, 1963b, 1979; Pennington, 1978), while the studies of van Someren & Rogers (van Someren, 1936, 1939a; Rogers & van Someren, 1925; van Someren & Rogers, 1925, 1926a, b, 1927, 1928a, b, 1929, 1931, 1932), supplemented by D.G. Sevastopulo’s volumes (unpublished) in the BMNH library, give a broad, if dated, view of the East African fauna. For the butterflies of the Indian Ocean islands, including Madagascar, the works of Manders (1908), Paulian, (1951b, 1956), Paulian & Viette (1968) and Desegaulx de Nolet (1982) all present useful details.

Compilations of available information on early stage biology appear in several group-based works. Aphytophagy (including myrmecophily) in the Lycaenidae has provided fertile ground for several reviews (Hinton, 1951; Henning, 1983b; Cottrell, 1984; Fiedler, 1991), each drawing freely on the classic works of lycaenid biology (Lamborn, 1914; Farquharson, 1922; Pomeroy, 1924; Clark & Dickson, 1971), themselves supplemented by a considerable volume of literature including Jackson (1937, 1947), Stempffer, (1957b), Cottrell (1965, 1981), van Someren (1974b), Heath (1983, 1987), Ackery (1990) and Henning & Henning (1988) (see also Callaghan, 1992). Other butterfly groups that have received particular attention include the Papilionidae (Villiers, 1957b), the Charaxinae (van Someren & van Someren, 1926; Henning, 1977; Darge, 1983; Smiles, 1985; Henning, 1989), the Danainae (Ackery & Vane-Wright, 1984), Hypolimnas (Vane-Wright et al., 1977), Junonia (Fontaine, 1985a) and Euphaedra (Hecq & Vuattoux, 1981). Of particular note is a study on Pseudopontia paradoxa (Heath, 1977), a taxonomically isolated species in the Afrotropical fauna.

Thus, in comparison to many other lepidopteran groups, information on the butterflies is abundant, if patchy. Attention has frequently been given to larger, more showy groups (various Nymphalidae, Pieridae, Papilionidae) or those known to be interesting (Lycaenidae), to the detriment of Hesperiidae and Satyrinae. Much has recently been accomplished in restoring a balance between available data for West Africa compared to the south and east. Perhaps the fauna of Madagascar, with so many endemic groups, represents the most important gap in current knowledge, although current research (by Claire Kremen and collaborators, Xerces Society) may soon rectify this situation.

GAZETTEER AND LIST OF REGIONAL WORKS

For the Afrotropical Region as a whole, the basic general works for the identification of butterflies are Aurivillius (1898-9), Aurivillius in Seitz (1908-1925), W.H. Evans (1937, 1947, 1956: Hesperiidae only), and D’Abrera (1980: Papilionoidea only). Peters’ (1952) checklist is the predecessor of the present work. A few families and subfamilies have been the subject of relatively recent monographs (e.g. Lycaenidae [to genus level]: Stempffer 1967b; Danainae: Ackery & Vane-Wright 1984; Acraeinae: Pierre 1984; Charaxinae: Henning 1989). Some works deal with major subregions (e.g. Pennington 1978, covers the whole of southern Africa; Villiers 1957, Stempffer 1957b, Boorman 1970, relate to much of west Africa; papers by Carcasson 1960, 1961b, 1963, cover certain families for east Africa). The only modern pocket guides for the whole region are Williams (1969) and Carcasson (1981). Biogeographic overviews based on butterflies are provided for Africa by Carcasson (1964c, an edited version of this paper is included in the present work – see Biogeography), for southern Africa by Cottrell (1978, 1985) and Pinhey (1978), the Arabian Peninsula by Larsen (1977, 1980, 1982, 1983a, 1984), Ethiopia by Carpenter (1935c), Madagascar by Corbet (1948), the African islands (by Plantrou & Blandin, 1987), the Palaearctic/Afrotropical boundary in the Sahel region by Bernardi (1962; see also Bernardi, 1989), and the montane forests of eastern Africa by Congdon & dejong (1992).

There are many national and local works, of varying vintage and quality, including various regional lists and similar accounts. In this section we attempt to give a guideline to the most useful faunistic works for each country and island. Once identification to genus level has been achieved, the specialised taxonomic literature referred to within the species accounts of the catalogue will often give access to additional information. Undoubtedly we have overlooked many valuable accounts, especially those in regional and non-English journals. None the less, even though far from complete, we hope this list will be of assistance in documenting and identifying local Afrotropical butterfly faunas. However, it should be recognised that all these works inevitably contain problems of synonymy and misidentification, many of which may not be evident at first sight, and most are also far from comprehensive, so that caution is always needed in their application.

Countries and islands are listed alphabetically, based on the gazetteer of Broadley & Minshull (1986), and with an indication of status and location (e.g. country of west Africa; island in Atlantic Ocean, etc.). Entries in bold capitals (modern countries and some islands) carry the references to local or regional works, with the most modern or comprehensive texts listed first, followed by older or more specialised references (works published before 1925 are not generally cited, except for some islands). Entries in non-bold capitals signify islands dealt with collectively elsewhere (each is cross referred). Entries in upper and lower case refer to regions which form part of modern states (cross referred in capitals), country and island synonyms (correct name in capitals), or marginal regions (such as many of the countries of north Africa) not considered to be part of the Afrotropical Region.

ABD AL KURI (island at mouth of Gulf of Aden: part of Yemen).

Ogilvie-Grant (1903), Rebel (1907).

Abyssinia (= ETHIOPIA).

Afars & Issas (= DJIBOUTI).

AGALEGA (islands east of northern Madagascar, Indian Ocean: part of MAURITIUS).

ALDABRA (islands at northern end of Mozambique Channel, Indian Ocean: administratively part of Seychelles; includes Aldabra, Picard, Assumption, Astove and Cosmoledo Archipelago).

Legrand (1965). See also Aurivillius (1909a), Fryer (1912), Cogan et al. (1971), Desegaulx de Nolet (1982).

Algeria (country of north Africa: part of Palaearctic Region).

ALPHONSE (island: part of Amirante group, SEYCHELLES).

AMIRANTE ISLANDS (part of SEYCHELLES).

Anglo-Egyptian Sudan (= SUDAN).

ANGOLA (country of south-western Africa).

No general account. See Bacelar (1948, 1956, 1958), Bivar de Sousa (1980), Bivar de Sousa & Fernandes (1964, 1966), Ladeiro (1956), Monard (1956).

ANJOUAN (island: part of COMORO ISLANDS).

ANNOBON (island: part of EQUATORIAL GUINEA).

Arabian Peninsula (mixed Afrotropical and Palaearctic fauna: see SAUDI ARABIA, TRUCIAL STATES, OMAN, DHOFAR, and YEMEN).

ARIDE (island: part of SEYCHELLES).

ASCENSION (island in Atlantic Ocean: tropical waif fauna).

Robinson & Kirke (1990).

ASSUMPTION (island: see ALDABRA).

ASTOVE (island: see ALDABRA).

Azores (islands in Atlantic Ocean: Palaearctic Region).

Basutoland (= LESOTHO).

Bechuanaland Protectorate (= BOTSWANA).

Belgian Congo (= ZAIRE).

Benguela (= part of ANGOLA).

BENIN (country of west Africa).

No general account. See Bacelar (1948), Condamin (1952), Stempffer (1952b), Bernardi (1952b).

BIJAGOS (islands, west Africa: part of GUINEA-BISSAU).

BIOKO (island: part of EQUATORIAL GUINEA).

Bophuthatswana (= part of SOUTH AFRICA).

BOTSWANA (country of southern Africa).

Larsen (in prep.), Pennington (1978), Pinhey (1968-76). See also van Son (1936b).

Bourbon (= RÉUNION).

British Bechuanaland (= part of SOUTH AFRICA).

British Cameroons (= part of CAMEROON and part of NIGERIA).

British Central Africa (= MALAWI).

British East Africa (= KENYA plus UGANDA).

British Kaffraria (= eastern cape province, part of SOUTH AFRICA).

British Somaliland (= part of SOMALIA).

BURKINA FASO (country of west Africa; formerly part of French West Africa).

No general account. See Talbot (1929c), and Hawker-Smith in Talbot (1929c).

BURUNDI (country of east Africa).

No general account. See Berger (1956), Fontaine (1988c, 1988e).

Cabinda (= part of ANGOLA).

Cabo Verde (= CAPE VERDE ISLANDS).

Caffraria (= part of SOUTH AFRICA).

CAMEROON (country of west Africa).

No general account. See Condamin (1953, 1958a, 1960), Bernardi (1955), Birket-Smith (1960), Darge (1983), Libert (1992).

Cameroun (= CAMEROON).

Canary Islands (Atlantic Ocean: part of Spain, Palaearctic Region, with very minor African influence).

Cape Colony (= part of SOUTH AFRICA).

CAPE VERDE (islands, Atlantic Ocean, including S. Antao, S. Vicente, S. Lucia, S. Nicolau, Sal, Boavista, Maio, S. Tiago, Fogo and Brava; predominantly Afrotropical fauna).

Nyström (1958), Riley (1968). See also Bacelar (1948), Traub & Bauer (1982).

Caprivi Strip (= part of NAMIBIA).

CARGADOS CARAJOS ISLANDS (part of SEYCHELLES).

Central African Empire (= CENTRAL AFRICAN REPUBLIC).

Central African Federation (= ZAMBIA plus ZIMBABWE plus MALAWI).

CENTRAL AFRICAN REPUBLIC (country of central Africa).

No general account. See Condamin & Soltani (1981).

CERF (island: part of SEYCHELLES proper, and another island by same name in Farquhar group, SEYCHELLES).

CHAD (country of northern central Africa).

No general account. See Bernardi (1962, 1964b).

Ciskei (see SOUTH AFRICA).

COETIVY (island: part of SEYCHELLES).

COMORO ISLANDS (Mozambique Channel, Indian Ocean; includes Grand Comore, Anjouan, Moheli and Mayotte).

Aurivillius (1909a); Strand (1917). See also Desegaulx de Nolet (1982), Turlin (1985).

CONGO (country of west Africa).

Bampton, Collins & Dowsett (1991). See also Condamin (1965c), Stempffer (1965), Descarpentries & Villiers (1966), Bernardi (1967), Berger (1967a), Fox (1968).

Congo-Belge (= ZAIRE).

Congo-Brazzaville (= CONGO).

Congo Free State (= ZAIRE).

Congo-Kinshasa (= ZAIRE).

Congo-Leopoldville (= ZAIRE).

COSMOLEDO (archipelago, including Menai, Moustique and Wizard: see ALDABRA).

Côte d’Ivoire (= IVORY COAST).

Côte d’Or (= GHANA).

Côte Française des Somalis (= DJIBOUTI).

COUSIN (islands: part of SEYCHELLES).

CURIEUSE (island: part of SEYCHELLES).

Dahomey (= BENIN).

Darfur (= part of SUDAN).

DESROCHES (island: part of Amirante group, SEYCHELLES).

Dhofar (part of OMAN).

DJIBOUTI (country of north-eastern Africa).

Genre (in Rougeot, 1984: 70-80).

EAGLE (island: part of Amirante group, SEYCHELLES).

EGYPT (country of north Africa: mixed Afrotropical and (mainly) Palaearctic fauna).

Larsen (1990).

EQUATORIAL GUINEA (country of west Africa; includes Rio Muni (mainland) and the islands of Annobon and Bioko in Gulf of Guinea).

Viejo (1984). See also Bacelar (1948), Bernardi (1953b), Beaurain (1985).

Eritrea (dealt with here as part of ETHIOPIA).

ETHIOPIA (country of north-eastern Africa; includes Eritrea).

Carpenter (1935c). See also Ungemach (1932), Gabriel (1949), Rougeot (1983, 1984).

EUROPA (island in Mozambique Channel, Indian Ocean).

No general account. See Aurivillius (1909a), Paulian (1956).

FARQUHAR (island group: part of SEYCHELLES).

Federation of Rhodesia & Nyasaland (included MALAWI, ZAMBIA and ZIMBABWE).

FÉLICITÉ (island: part of SEYCHELLES).

Fernando Poo (= Bioko island, part of EQUATORIAL GUINEA).

French Cameroons (= CAMEROON).

French Equatorial Africa (included CHAD, CENTRAL AFRICAN REPUBLIC, GABON and CONGO).

French Guinea (= GUINEA).

French Somali Coast (= DJIBOUTI).

French Soudan (= MALI).

French Territory of the Afars and Issas (= DJIBOUTI).

French West Africa (included MAURITANIA, SENEGAL, GUINEA, MALI, IVORY COAST, BURKINA FASO, BENIN and NIGER).

FRIGATE (island: part of SEYCHELLES).

GABON (country of west Africa).

No general account. See Stempffer (1970b).

Gaboon (= GABON).

GAMBIA (country of west Africa).

No general account. See Gillies (1985); also SENEGAL.

Gazankulu (= part of SOUTH AFRICA).

GHANA (country of west Africa).

No general account. See Usher (1979, and MS [1985] b).

Glorieuses – see ILES GLORIEUSES.

Gold Coast (= GHANA).

GRAND COMORE (island: part of COMORO ISLANDS).

Guinea. In older literature, the west coast of Africa, roughly from Senegal to Walvis Bay.

GUINEA (country of west Africa; formerly French Guinea).

No single modern comprehensive account. See De Fleury (1926), Stempffer (1953b, 1954e, 1963b), Bernardi (1954), Evans (1954), Berger (1962), Bernardi & Condamin (1963), Condamin (1963c), Condamin & Roy (1963).

GUINEA-BISSAU (country of west Africa; formerly Portuguese Guinea; includes Bijagos Archipelago).

Bivar de Sousa & Passos de Carvalho (1987). See also Bacelar (1948, 1949), Merten (MS, 1987).

Guinea Española (= EQUATORIAL GUINEA).

Guineé Française (= GUINEA).

Haute Volta (= BURKINA FASO).

Ile de Bourbon (= RÉUNION).

Ile de France (= MAURITIUS).

Ile Maurice (= MAURITIUS).

ILE SAINTE MARIE (island: part of MALAGASY REPUBLIC).

ILES GLORIEUSES (islands in Indian Ocean, north of Madagascar).

No general account. See Paulian (1956).

Italian East Africa (included Eritrea, ETHIOPIA and part of SOMALIA).

Italian Somaliland (= part of SOMALIA).

IVORY COAST (country of west Africa).

No general account. See Riley (1926b), Berger (1962), Stempffer (1966, 1969b), Condamin & Vuattoux (1973), Blandin, Vuattoux & Plantrou (1976), Vuattoux & Blandin (1977, 1979), Pierre & Vuattoux (1978).

JUAN DE NOVA (island in Mozambique Channel, Indian Ocean: part of MALAGASY REPUBLIC).

No general account. See Aurivillius (1909a).

Kaffraria (= part of SOUTH AFRICA).

Kamerun (= CAMEROON plus part of NIGERIA).

Kangwane (= part of SOUTH AFRICA).

Katanga (= part of ZAIRE).

KENYA (country of east Africa).

Larsen (1991). See also Rogers & van Someren (1925), van Someren & Rogers (1925, 1926a, 1926b, 1927, 1928a, 1928b, 1929, 1931, 1932), van Someren (1935, 1936, 1939a, 1939b), Wiley & Hudson (1943), R. T. Evans (1944, 1946), Stoneham (1951-65), Sevastopulo (1973-4).

Kivu (= part of ZAIRE).

Kwandebele (= part of SOUTH AFRICA).

Kwazulu (= part of SOUTH AFRICA).

LA DIGUE (island: part of SEYCHELLES).

Lebowa (= part of SOUTH AFRICA).

LESOTHO (see SOUTH AFRICA).

LIBERIA (country of west Africa).

Fox et al. (1965). See also Picard (1950), Stempffer (1950c), Condamin (1951), Berger (1954d), Stempffer & Bennett (1956a).

Libya (country of north Africa; mainly Palaearctic fauna).

Macias Nguema Biyogo (= BIOKO, part of EQUATORIAL GUINEA).

Madagascar (= MALAGASY REPUBLIC).

Madeira (island in Atlantic Ocean; Palaearctic Region).

MAFIA (island group, including Mafia and Songo, in Mozambique Channel, Indian Ocean: part of TANZANIA).

No general account. See Aurivillius (1909a).

MAHÉ (island: part of SEYCHELLES).

MALAGASY REPUBLIC (island country, separated from east Africa by the Mozambique Channel).

Paulian (1951b). See also Viette (1956, 1971b), Paulian (1956), Paulian & Viette (1968).

MALAWI (country of east Africa).

Gifford (1965). See also Beaurain (1989).

MALI (country of north-western Africa).

No account.

MANDA (island: part of KENYA).

No general account. See Aurivillius (1909a).

Mascarene Islands (Indian Ocean: see MAURITIUS, RÉUNION, and RODRIGUES).

MAURITANIA (country of north-western Africa).

No general account. See Bernardi (1966c).

MAURITIUS (island in Indian Ocean, east of Madagascar; includes Round Island, now apparently devoid of butterflies).

Williams (1989). See also Manders (1908), Vinson (1938), Desegaulx de Nolet (1982), Barnes (MS, 1982).

MAYOTTE (island: part of COMORO ISLANDS).

MBINI (island: part of EQUATORIAL GUINEA).

MOHELI (island: part of COMORO ISLANDS).

Morocco (country of north Africa; Palaearctic fauna).

Moyen Congo (= CONGO).

MOZAMBIQUE (country of southern Africa).

Pennington (1978). See also Bacelar (1948), Pinhey (1976).

NAMIBIA (country of southern Africa).

Pennington (1978).

NIGER (country of north-western central Africa).

No general account. See Bernardi (1950), Villiers (1950), Stempffer (1950d).

NIGERIA (country of west Africa).

Cornes (1969, 1971), Comes, Riley & St Leger (1972). See also Boorman & Roche (1957, 1959), Boorman (1961, 1965, 1973), St Leger (1965, 1984, 1989), Hopkins (1970), Riley & Cornes (1970, 1971), Cornes & Riley (1972), Medler (1980), Larsen, Riley & Cornes (1981), Dowsett, Hecq & Knoop (1989), Hitchings, Knowles & Knowles (MS, 1990).

NORTH (island: part of SEYCHELLES).

North Yemen (= part of YEMEN).

Northern Rhodesia (= ZAMBIA).

NOSSI-BÉ (island off northwest coast of Madagascar: part of MALAGASY REPUBLIC).

Nyasaland (= MALAWI).

OMAN (state in Arabian Peninsula: Palaearctic and Afrotropical fauna).

Larsen & Larsen (1980). See also Larsen (1977, 1983, 1984).

Oubangui-Chari (= CENTRAL AFRICAN REPUBLIC).

Pagalu (= ANNOBON, part of EQUATORIAL GUINEA).

PATTA (coastal island north of mouth of Tana River: part of KENYA).

No general account. See Aurivillius (1909a).

PEMBA (island north of Zanzibar: part of TANZANIA).

Kielland (1985d). See also Aurivillius (1909a), Kielland (1990b).

People’s Democratic Republic of Yemen (= part of YEMEN).

Pigalu (= ANNOBON, part of EQUATORIAL GUINEA).

PLATTE (island: part of SEYCHELLES).

Portuguese Congo (= Cabinda, part of ANGOLA).

Portuguese East Africa (= MOZAMBIQUE).

Portuguese Guinea (= GUINEA-BISSAU).

PRASLIN (island: part of SEYCHELLES).

PRINCIPE (island: see SAO TOMÉ & PRINCIPE, part of EQUATORIAL GUINEA).

PROVIDENCE (island: part of SEYCHELLES).

Qwaqwa (= part of SOUTH AFRICA).

RECIF (island: part of SEYCHELLES).

Republic of South Africa (= SOUTH AFRICA).

RÉUNION (island in Indian Ocean, east of Madagascar).

Desegaulx de Nolet (1982). See also Manders (1908), Vinson (1938).

Rhodesia (= ZIMBABWE, or ZAMBIA plus ZIMBABWE (1899-1911)).

Rio del Oro (= SAHRAWI REPUBLIC).

Rio Muni (= mainland part of EQUATORIAL GUINEA).

RODRIGUES (island in Indian Ocean, east of Madagascar),

Desegaulx de Nolet (1982). See also Snell & Tams (1920), Vinson (1938).

Rodriguez (= RODRIGUES).

ROUND (island; part of MAURITIUS).

Ruanda (= RWANDA).

Ruanda-Urundi (= RWANDA plus BURUNDI).

RWANDA (country of east Africa).

No general account. See Berger (1956), Turlin (1979, 1980), Fontaine (1988a, 1988c), Monfort (1987, 1989), Joris-Mayence & Monfort (1989, 1990).

Sahara Español (= SAHRAWI REPUBLIC).

SAHRAWI REPUBLIC (country of north-western Africa).

No general account. See Bernardi (1966c).

ST BRANDON (= Cargados Carajos islands, part of SEYCHELLES).

ST HELENA (island in Atlantic Ocean; Wollaston lists four widespread species, all of which occur in Africa).

Wollaston (1879).

St Johanna (= ANJOUAN, part of COMORO ISLANDS).

ST PIERRE (island: part of SEYCHELLES).

ST MARIE (island: part of MALAGASY REPUBLIC, off north-east coast of Madagascar).

No general account. See Aurivillius (1909a).

SÃO TOMÉ & PRINCIPE (islands in Gulf of Guinea, Atlantic Ocean; part of EQUATORIAL GUINEA).

Viejo (1984). See also Bacelar (1948), Pyrcz (1992), Riley (1928a).

SAUDI ARABIA (country of Arabian Peninsula; Palaearctic and Afrotropical fauna).

Larsen (1983, 1984). See also Gabriel (1954), Pittaway (1985).

SENEGAL (country of west Africa).

No general account. See Condamin (1957, 1962, 1964, 1969, 1990), Stempffer (1963a), Bernardi & Condamin (1964), Berger (1968).

Senegambia (proposed union of SENEGAL and the GAMBIA).

Senegambia-Niger (= MALI).

SEYCHELLES (country and group of islands in Indian Ocean; includes ALDABRA group (q.v.), together with the Amirante Islands, Cargados Carajos Islands, Coetivy, Farquhar Islands, Platte, Providence and St Pierre; the Seychelles proper includes Mahé, Cerf, Silhouette, North, Praslin, Aride, Cousin Islands, Curieuse, Félicité, Frigate, La Digue, Recif and The Sisters).

Legrand (1965). See also Fryer (1912), Cogan et al. (1971), Desegaulx de Nolet (1982).

SIERRA LEONE (country of west Africa).

Belcastro (1986a, 1990b). See also Condamin (1971), Owen & Owen (1972, 1974).

SILHOUETTE (island: part of SEYCHELLES).

SISTERS, The (island: part of SEYCHELLES).

SOCOTRA (island at mouth of Gulf of Aden: part of Yemen).

Ogilvie-Grant (1903), Rebel (1907).

SOMALIA (country of north-eastern Africa).

Talbot (1932c). See also Butler (1886c).

SONGO (island: part of TANZANIA; see MAFIA).

Soudan Français (= MALI).

SOUTH AFRICA (including the independent states of LESOTHO and SWAZILAND, and the homelands Bophuthatswana, Ciskei, Gazankulu, Kangwane, Kwandebele, Kwazulu, Lebowa, Qwaqwa, Transkei and Venda).

Pennington (1978; at time of writing a second edition is near completion). See also Murray (1935), van Son (1949a, 1955, 1963a, 1979), Swanepoel (1953a), Pennington (1963), Pinhey (1965), Kloppers & van Son (1978), Claassens & Dickson (1980), Vari & Kroon (1986), Migdoll (1988), Henning & Henning (1989b).

South West Africa (= NAMIBIA).

South Yemen (= part of YEMEN).

Southern Cameroons (= part of NIGERIA).

Southern Nigeria (= part of NIGERIA).

Southern Rhodesia (= ZIMBABWE).

Spanish Guinea (= EQUATORIAL GUINEA).

Spanish Sahara (= SAHRAWI REPUBLIC).

Spanish West Africa (= SAHRAWI REPUBLIC plus part of Morocco).

SUDAN (country of north east Africa).

No general account. See Carpenter (1928, 1938), Wilson (1949, 1953).

SWAZILAND (see SOUTH AFRICA).

Tanganyika (= part of TANZANIA).

TANZANIA (country of east Africa).

Kielland (1990b). See also Aurivillius (1909a), Haldane (1969), Whybrow (1940), Carcasson (1966), Berger (1975, 1980), Stempffer (1976), Cordeiro (1990), Kielland (1968-9, 1978, 1979, 1980-81, 1985d).

Tchad (=CHAD).

TOGO (country of west Africa).

No general account. See Condamin (1952), Stempffer (1952b), Bernardi (1952b).

Transkei (= part of SOUTH AFRICA).

TROMELIN (island, east of northern Madagascar, Indian Ocean).

TRUCIAL STATES (state in Arabian Peninsula).

Larsen (1984).

Tunisia (country of north Africa; Palaearctic Region).

Ubangi-Chari (= CENTRAL AFRICAN REPUBLIC).

Ubangi-Shari-Chad (= CENTRAL AFRICAN REPUBLIC).

UGANDA (country of east Africa).

Rogers & van Someren (1925), van Someren & Rogers (1925, 1926a, 1926b, 1927, 1928a, 1928b, 1929, 1931, 1932), van Someren (1935, 1936, 1939a), R.T. Evans (1946). See also Carpenter (1934b), Gabriel (1939), Jackson (1956, 1961), Stempffer & Jackson (1962).

Union of South Africa (= SOUTH AFRICA).

Upper Senegal-Niger (= MALI plus BURKINA FASO).

Upper Volta (= BURKINA FASO).

Urundi (= BURUNDI).

Venda (= part of SOUTH AFRICA).

Walvis Bay (= part of NAMIBIA; formerly South African enclave within NAMIBIA).

Western Sahara (= SAHRAWI REPUBLIC).

YEMEN (state in Arabian Peninsula).

Larsen (1982a, 1984).

Yemen Arab Republic (= part of YEMEN).

ZAIRE (country of central Africa).

Berger (1981). See also Berger (1940, 1950, 1967b), Dufrane (1939, 1940, 1945, 1948a, 1953), Overlaet (1955), Mathot & Kadima (1971), Fontaine (1988a, 1988c, 1988e).

ZAMBIA (country of south-central Africa).

Neave (1910), Dening (MS, 1979). See also Pinhey & Loe (1977).

ZANZIBAR (island: part of TANZANIA).

ZIMBABWE (country of southern Africa).

Pennington (1978), Cooper (1973), Pinhey (1949b). See also Pinhey (1949a), Hancock (1982).

Zimbabwe-Rhodesia (= ZIMBABWE).

Zululand (= part of SOUTH AFRICA).

BIOGEOGRAPHY

This section is based directly on Bob Carcasson’s pioneering and influential publication, A preliminary survey of the zoogeography of African butterflies (Carcasson, 1964c). Bob’s first manuscript catalogue of African butterflies formed the basis of the 1964 paper. In that account he indicated that much of his information came from the great collections in London and Nairobi, acknowledging help from the late T.H.E. Jackson and V.G.L. van Someren, as well as staff of The Natural History Museum, London (BMNH). His desire to publish the catalogue, the database on which the preliminary survey was founded, later brought him into renewed contact with the BMNH, and led to the idea of the present work.

Some time after Bob’s death, the editors decided to include his original biogeographic essay as one of the introductory texts. We felt that this would be fitting, not only as a memorial, but also to make his important paper more readily available. However, on getting down to the task, we found ourselves faced with a number of practical problems. A facsimile reprinting within the catalogue would have looked out of place, and given no opportunity to update geographical or biological nomenclature, or correct a variety of minor errors evident to us, nor would it have solved certain difficulties with the figures. On the other hand, a major updating would have run the risk of distorting Bob’s firmly held and highly original ideas.

In the event we have opted for a compromise. The text has been updated with respect to geographical and scientific names, metres substituted for feet and so on, and a variety of minor errors corrected and punctuational idiosyncracies changed. Authorities for butterfly species-group names have been removed throughout, as these are all available in the catalogue. The text has been more or less lightly edited, notably by breaking up some of the longer sentences, editing out or adding a few words here and there, shifting some sections round, all with the intention of making the flow of ideas easier to appreciate.

Many of the numbers, such as apparently precise totals for species or genera within regions, were not based on explicit information available to us. As knowledge of the Afrotropical fauna has progressed, these undoubtedly needed updating. In a few places we have done so, in others we have simply rounded the figures up, but in most cases we have decided to omit them altogether. Likewise, we have also omitted all the original maps and, with one exception, the tables. The maps were of rather low quality and, in relation to the text, the loss of information seemed minimal. In a few places, where subsequent work known to the editors points to significantly different ideas or conclusions compared with Bob’s original text, editorial notes and references have been added [generally within square parentheses, thus]. References to many other relevant publications can also be found in the section on Regional Works.

Overall, we have been guided by the notion of minimally editing Bob’s published text, almost as if it was a posthumous work submitted to a mainstream biological journal. We have endeavoured to be true to the spirit, but not necessarily to the letter, number, word or sentence.

BUTTERFLIES AND THE AFROTROPICAL REGION

Having attracted a great deal of attention, butterflies are sufficiently well known to yield a vast amount of positive distributional information, as well as some negative evidence of unquestioned validity. Although very mobile, many species are restricted to particular habitats through their dependence on special food plants, and by the narrow compass of their climatic optima. Minor changes, genetic or otherwise, due to the segregation of populations or to the influence of climatic factors, are often expressed by conspicuous variations in colour and pattern. This combination of factors makes the butterflies an ideal group for the study of zoogeographical problems.

Carcasson (1964c) originally attempted to determine the dominant patterns of distribution of African butterflies, relate such patterns to the ecological potential of the environment, and assess the impact of past climatic changes on the distribution of the recent fauna. The whole of the Afrotropical Region (continental Africa south of the Sahara, Madagascar and adjacent islands, and southern Arabia) is considered. Mediterranean Africa is inhabited by a Palaearctic fauna with very few Afrotropical penetrations, and is therefore not included. The butterfly fauna of the Malagasy Republic is treated in toto and its relationship to that of continental Africa is examined, but the detailed geographic distribution of species within Madagascar is not discussed.

The butterfly fauna of the Afrotropical Region, as listed in the present catalogue, consists of 301 genera and of some 3607 known species. It is likely that at least 90% of the total species have now been described, although the distribution ranges of many of the rarer ones remain very imperfectly known. A good many Lycaenidae probably still await discovery and description, particularly in the equatorial forest region, and also some of the Satyrinae in the remoter mountains of eastern and southern Africa [and possibly Madagascar — eds]. Critical revisions of the larger and more difficult groups, especially in the Nymphalidae and Lycaenidae, will no doubt yield considerable additions to the total number of Afrotropical species. Nearly 98% of species, and 76% of butterfly genera in the present list do not occur outside the Region [see additional discussion, pp 42-47 — eds].

AFRICAN REGIONAL GEOGRAPHY

The African continent is shaped rather like an irregular pear, with the blunt end forming the Mediterranean shore, and the opposite end marking the junction of the south Atlantic and Indian oceans. The northern and southern extremities of the continent are placed near the 35th parallel in their respective hemispheres and the equator passes through the centre of Africa, imparting a remarkable symmetry of climate and vegetation to the continent. This symmetry is upset, however, by the fact that the northern part of Africa occupies a far greater area than the southern, and by the presence of a high, irreglar plateau in the east, from Ethiopia to South Africa and with a westerly extension stretching across Zambia and southern Zaire to Angola. The eastern plateau is broken by a depression in north Kenya and north Uganda, which separates the Abyssinian highlands in the north from the main plateau in the south. In Zimbabwe, Shaba (Zaire) and Angola the plateau is very uniform, forming part of an ancient peneplain and consisting of old crystalline rocks. Further north it has been disturbed by extensive rifting and by much recent volcanic activity. The central part of the continent is characterised by the presence of a vast shallow depression (Congo Basin) which is contained by the plateau of the great lakes in the east, by the Angolan highlands in the south and by the Congo-Nile watershed and the Cameroon highlands in the north. There are also isolated high massifs in the Sahara (Ahaggar and Tibesti) and in West Africa (Fouta Djalon).

The large island of Madagascar (Malagasy Republic) is situated in the Indian Ocean, some 400km from the African mainland at its nearest point, and stretches from 13°S. to 26°S. Madagascar has a central ridge running from north to south, reaching 2600m at some points. There are several groups of small islands in the Indian Ocean, such as Réunion, Mauritius, Rodrigues, the Comoros, the Aldabra group and the Seychelles, and several in the gulf of Guinea (Bioko, Principe, Sāo Tomé and Annobon). [Corbet (1948) and Plantrou & Blandin (1987) have reviewed the butterfly faunas of Madagascar and the smaller islands, respectively — eds.]

AFRICAN CLIMATE AND VEGETATION

The equatorial belt of Africa, from the Atlantic to the western edge of the eastern plateau (30°E.) and from approximately 5°N. to 5°S. enjoys a typically equatorial climate, with high mean temperatures all the year round and well distributed rainfall. This area is naturally occupied by an almost unbroken block of typical evergreen equatorial forest which continues along the Guinea coast as far as Conakry (9°N.). This forest is divided into two parts by a broad band of savanna which reaches the coast in Togo and Benin. As one moves away from the equator, north or south, the rainfall gradually decreases, the annual dry seasons become longer, and seasonal and daily temperature oscillations become increasingly pronounced, until one encounters conditions of extreme aridity at about 20° latitude north and south. With the gradual desiccation the equatorial forest gives way to deciduous woodland of decreasing density, then to wooded steppe, open steppe, arid scrub and eventually desert. Beyond the desert areas there is a moderate winter rainfall together with a well defined cool season, giving rise to a vegetation of evergreen shrubs and trees with small, leathery leaves: the Mediterranean flora of North Africa, and the analogous but completely unrelated Cape maquis of South Africa.

In the northern hemisphere, the desert regions (Sahara) occupy a much greater area than in the southern dry zone (Kalahari), where extreme desert conditions are limited to a narrow coastal strip in Namibia and southern Angola (the Namib). This is due to the much greater land area of the northern part of Africa.

The extension of continental climatic influences in the north is also responsible for the much more rapid transition from equatorial forest to desert. In the south this transition is slower and less regular, due no doubt to the greater prevalence of maritime climatic influences and to the great south-central plateau, with its almost uniform cover of deciduous woodland. The southern transition from desert (Kalahari) to Mediterranean type vegetation (Cape maquis) includes a large area with a very characteristic vegetation of low-growing succulent plants (Karroo), which lacks any counterpart in the northern hemisphere.

In the eastern part of Africa the typical succession of vegetation is strongly modified by the plateau. Equatorial conditions are limited to a narrow coastal belt which, however, extends further south than on the west coast, due to the effect of the warm Mozambique current and the south-east monsoon. In this narrow belt there are many small relics of equatorial forest, isolated from one another and forming a complex mosaic with deciduous woodlands and savanna. These forests frequently extend to the eastern escarpment of the plateau, where the moist winds from the Indian Ocean are forced to condense their water content. Beyond the escarpment lies a vast table-land consisting of savanna and steppe, with scattered areas of highland forest at higher elevations, and subalpine grassland and moorland on the higher mountains, above the tree line. It is interesting to note that the highland forest, though confined to scattered mountains, is surprisingly uniform in botanical composition and occurs from northern Ethiopia to South Africa, but at progressively lower altitudes as one proceeds further from the equator.

The Knysna forest in the Cape, though situated at sea level, is similar to forest areas occurring in Kenya and Uganda above 1800m. The very few scattered surviving montane forests of West Africa (Cameroon and Angola) are also similar to their east African counterparts. The mountains in Liberia, Sierra Leone and Guinea are either too low to have developed true montane forest, or have been so badly denuded by man that it is difficult to ascertain the true nature of their autochthonous vegetation.

In Madagascar rainfall decreases progressively from east to west. Most of the east coast is naturally covered by a broad belt of moist evergeen lowland forest which gives way to grass savanna and grass steppe in the centre, with scattered relics of montane forest at higher elevations. The west coast, facing the continent of Africa, is largely covered by xerophilous thicket and scrub.

CLIMATIC CHANGES DURING LATE TERTIARY AND QUATERNARY

According to Joly (1930) and Du Toit (1939) the general outline of the African continent has changed little since early Cretaceous times. According to Krenkel (1925), Madagascar became detached from Africa near the end of the Miocene period (about 15 million years ago). There are also indications that, in the past, Africa may have been more closely connected with Arabia and India (Moreau, 1952). [Note: Carcasson wrote his original version of this section without access to modem knowledge of plate tectonics, etc.; atlases of continental displacement, such as Owen (1983), suggest afar older separation of Madagascar from Africa, at 150-200 mya, and that Madagascar remained in substantial contact with southern India for a long while after its rift from the African mainland, up to about 100 mya — eds.]

Available evidence suggests that most of Africa during the early Miocene was a gently eroded, uniform peneplain. Later in the Miocene the Abyssinian and East African plateaux were uplifted. This process intensified during the Pliocene period, during which there was also a great deal of tectonic rifting, accompanied by much volcanic activity, and by the downwarping of the Congo Basin (which became a large lake). During the Pliocene most of the large volcanic masses began to build up (including Mt Cameroon). Large igneous blocks such as Ruwenzori and possibly the Usambara Mountains were uplifted at this time. Volcanic activity was intensified during the subsequent Pleistocene period and indeed persists to the present day in some areas.

There are strong reasons for believing that up to the mid-Miocene the world enjoyed a generally mild climate, with an extension of the tropical belt to some 55°N. latitude and 40°S. Outside these limits the climate was probably temperate, except in the immediate vicinity of the poles (Moreau, 1952).

The equatorial forest belt appears to have been subjected to periodic contractions and expansions, but to have maintained a permanent and stable core since Mesozoic times. However, the fact that the African equatorial forest is poor in families typical of the tropics, such as palms, orchids, bamboos and aroids, and its low density of plant species (25 000 compared to 40 000 in South America: Good, 1952) suggests that it may have contracted to small areas during periods of extreme aridity. Aubréville (1949) thinks that during Tertiary times the equator was a great deal further north and that it has moved to its present position during the Pliocene and Pleistocene; this would account for the astonishing richness of the Cape flora, which although confined to a very small area at present, probably evolved in an area of far greater extent [Owen’s (1983) maps suggest that the equator was only substantially further north sometime before the beginning of the Tertiary, at about 90 mya — eds]. Aubréville also believes that the poverty of species of the northern deciduous woodland, compared to the homologous southern deciduous woodland, is due to the fact that only a few species managed to squeeze past between the forest and the plateau when the equator moved south. The scarcity of floral exchanges between tropical Africa and Mediterranean Africa suggests that, whatever its past position, the Sahara has been an effective barrier for a very long time, possibly since the mid-Miocene.

The series of tectonic movements and climatic oscillations which began in the late Miocene and gathered strength in the Pliocene reached a climax in the Pleistocene. During this period the northern hemisphere underwent a series of major glaciations, corresponding with similar though less extensive glaciations in the southern hemisphere. During periods of glaciation at higher latitudes, the tropical belt appears to have enjoyed a cooler and wetter climate and in Africa there are indications of at least three such pluvial periods, during which the forest covered a greater area than at present.

On the other hand the interglacial maxima, and particularly the long Mindel-Riss interglacial, were milder at high latitudes than the present climate, but hot and arid in the tropics (interpluvials). In Africa most of the shallower lakes dried out during the interpluvial phases and the forest contracted, possibly surviving only in moister areas such as upper Guinea and Cameroon, and in the vicinity of mountains such as Ruwenzori, Elgon and the Usambaras. Lake Nyasa and Lake Tanganyika, being very deep, persisted and had time to evolve rich endemic faunas (Charlesworth, 1957).

SPECIATION, SUBSPECIATION AND VICARIANCE

Individuals which interbreed freely under natural conditions are said to be members of the same species. When the range of a species is broken (disjunction) by climatic changes or tectonic movements, or by any other circumstances, the resulting populations may, in isolation from one another, develop different characters but still retain the ability to interbreed with one another, should their ranges coalesce again. Such populations are usually called subspecies or races, and if a subspecies is isolated from its parent stock long enough to lose its ability to interbreed with it, then it has become a full species.

This process is going on continually, and species are breaking up to form subspecies which may coalesce again before full speciation has taken place, or may develop into full species if isolation continues. Between these conditions there is every possible transition in status, but such intermediate stages cannot be expressed in terms of conventional nomenclature. Even full species differ in rank, some being older and more isolated than others. All taxonomic groupings, however natural the system of classification, have an element of artificiality, since the only absolute unit is the individual, every individual being unique.

When two closely related populations are segregated from one another by geographical barriers, but occupy similar habitats, they are called geographical vicariants. When two such populations are segregated by their preference for different habitats, they are called ecological vicariants. Geographic vicariance can be at the specific or subspecific level; ecological vicariance, which may have been originally caused by geographic segregation, is nearly always specific since it often involves a degree of overlapping (sympatrism) which would lead to interbreeding and integration if the populations concerned were not specifically distinct. Geographic vicariants do not normally overlap and are therefore called allopatric. Vicariance is well illustrated by the following examples:

GEOGRAPHICAL VICARIANTS

Papilio menestheus/P. lormieri/P. ophidicephalus. All three are forest species and very closely related. Papilio menestheus occurs from Senegal to Cameroon; P. lormieri from Nigeria to Lake Tanganyika and to the Rift Valley in west Kenya; P. ophidicephalus from eastern Kenya through eastern and southern Tanzania to Malawi, Zimbabwe, Mozambique and South Africa. Both P. lormieri and P. ophidicephalus have evolved several distinct subspecies, especially ophidicephalus in South Africa, where its forest habitat is reduced to isolated relics.

Papilio hesperus/P. horribilis/P. pelodorus. All three are very closely related forest species. Papilio hesperus occurs from Ivory Coast to southern Sudan, Zambia, Zaire and Angola; P. horribilis is found in the upper Guinea forest block, from Ghana to Sierra Leone; P. pelodorus occurs in Malawi, Mozambique, Tanzania and Zambia. In Zambia the two eastern species are separated by the narrow Luangwa Rift, hesperus occurring at Mpika (R.H. Carcasson, May 1953) and pelodorus in the Nyika plateau (C.B. Cottrell, in litt. to Carcasson).

Aterica galene/A. rabena. Aterica galene is common in lowland forest throughout Africa; A. rabena is found in Madagascan forests. The two species, which mimic different models and are therefore very different in superficial appearance, are very closely related (they are the only representatives of their genus).

ECOLOGICAL VICARIANTS

Charaxes varanes/C. fulvescens/C. acuminatus. These three species are closely related, and very distinct from all other members of the genus, but are replaced by geographical vicariants in Arabia, Socotra and the Comoros. C. varanes occurs in semi-arid scrub, savanna and deciduous forest throughout Africa, but hardly ever in evergreen forest. C. fulvescens is common in evergreen forest from sea level to about 1500m from Senegal and Sierra