Biodiversity of Angola Brian J. Huntley · Vladimir Russo Fernanda Lages · Nuno Ferrand Editors Science & Conservation: A Modern Synthesis Foreword by His Excellency President of the Republic of Angola João Manuel Gonçalves Lourenço Biodiversity of Angola Brian J. Huntley • Vladimir Russo Fernanda Lages • Nuno Ferrand Editors Biodiversity of Angola Science & Conservation: A Modern Synthesis UNESCO Chair on Life on Land University of Porto Porto, Portugal United Nations Educational, Scientific and Cultural Organization ISBN 978-3-030-03082-7 ISBN 978-3-030-03083-4 (eBook) https://doi.org/10.1007/978-3-030-03083-4 Library of Congress Control Number: 2018966550 © The Editor(s) (if applicable) and The Author(s) 2019 Open Access This book is licensed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence and indicate if changes were made. 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Huntley CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos Universidade do Porto Vairão, Portugal Fernanda Lages ISCED – Instituto Superior de Ciências da Educação da Huíla Lubango, Angola Vladimir Russo Fundação Kissama Luanda, Angola Nuno Ferrand CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos Laboratório Associado, Campus de Vairão Universidade do Porto Vairão, Portugal Departamento de Biologia, Faculdade de Ciências Universidade do Porto Porto, Portugal Department of Zoology, Auckland Park University of Johannesburg Johannesburg, South Africa . This book is an open access publication To the new generation of Angolan students of biodiversity. May they stand on the shoulders of giants: the founders of Angola’s biodiversity science Friedrich Martin Josef Welwitsch (1806–1872) José Vicente Barbosa du Bocage (1823–1907) José Alberto de Oliveira Anchieta (1832–1897) Johannes (John) Gossweiler (1873–1952) Contributors to this volume record with sadness the passing of William Roy (Bill) Branch (1946–2018) - herpetologist, indefatigable field researcher and mentor of young Angolan scientists vii Foreword Angola occupies only four percent of the terrestrial area of the African continent, yet it possesses the highest number of biomes of any African country. It is second in terms of the number of ecoregions represented within its borders. It has ecosystems as diverse as the rainforests of Maiombe of Cabinda to the vegetation-less dunes of Namibe and the endless savannas and woodlands of the Cuando Cubango to the tiny remnant forests of the highest valleys of Mount Moco in Huambo. It is the only home to the most magnificent mammal in the world – the Giant Sable Antelope. It was in Angola that one of the most extraordinary plant species Welwitschia mirabilis was discovered and described – the enigmatic ‘living fossil’ of the desert. It even puzzled Charles Darwin, who compared its evolutionary importance in the plant kingdom to that of the Duck-billed Platypus in the animal kingdom. Considering real fossils, Angola’s history goes back hundreds of millions of years, to the earliest known living organisms, the bacterial stromatolites of the limestones His Excellency the President of the Republic of Angola, João Manuel Gonçalves Lourenço viii of Bembe and Humpata. Angola’s fossils range in size from microorganisms to the gigantic dinosaur – Angolatitan adamastor – recently discovered in the sediments of the Bengo coastline. Yet, despite this globally significant natural wealth, Angola remains one of the least well-documented countries in the world in terms of its bio- diversity. This situation is about to change. Angolan scientists have collaborated with over 40 colleagues from 7 countries to produce a new synthesis of knowledge of Angola’s remarkable biodiversity. They have produced a magnificent volume that seeks to review all what is known about Angola’s biodiversity, especially that which has been revealed through studies undertaken in the twenty-first century. For several decades, field studies were ren- dered nearly impossible because of the disruptions of war. But since peace was achieved in 2002, a new generation of research has been made possible, bringing many foreign specialists into partnerships with Angolan scientists and institutions and introducing new technologies that have helped stimulate an unprecedented wave of research activity. Angola’s indigenous knowledge acquired over millennia provided the founda- tions to the information documented and materials collected by visiting researchers from the eighteenth to twenty-first centuries. Many detailed accounts have been published over the past century and more, but often in scientific journals and official reports that have been lost with the passage of time. More important, the knowledge that does exist is fragmentary and largely inaccessible to Angolan students and researchers. Much is published in foreign languages, and thus the inexistence of a comprehensive synthesis of studies on Angola’s fauna, flora and ecosystems is a challenge for young researchers. Angolan students, researchers and government officials have very limited access to data sources where they can find a reliable, science-based and up-to-date summary of that which has been recorded on the country’s biodiversity. It was the need for an integrated ‘state-of-knowledge’ summary, recognised dur- ing the past decade by many Angolan university and government colleagues, that was the catalyst that stimulated this important project. From humble beginnings by a few Angolan and foreign partners, the effort expanded into the present volume of over 500 pages of authoritative accounts on our landscapes, seascapes, vegetation, flora and fauna, its past and future. Most importantly, this work identifies the exciting opportunities for research and conservation that Angolan scientists, conservationists, government officials and the general public can embrace as the country moves for- ward to an ever greater and more prosperous and environmentally sustainable future. It is a pleasure to endorse this valuable contribution to the new wave of Angolan scientific and conservation literature, a source of inspiration to our students and a reminder to all our leaders, young and old, of our responsibility to treasure and safeguard Angola’s unsurpassed, but vulnerable, biodiversity and natural resources. President of the Republic of Angola Luanda, Angola João Manuel Gonçalves Lourenço Foreword ix Acknowledgements This book was conceived as a collaborative and voluntary endeavour shared by students of Angola’s biodiversity. Contributors were drawn from Angola, Britain, Germany, Namibia, Portugal, Swaziland, South Africa, the Netherlands and the United States. To all contributors to this synthesis, the Editors offer their thanks for the unstinting efforts of the synthesis team to meet the strict demands of quality and of timelines. Over the past decades, biodiversity research in Angola has been encouraged by successive leaders within the government, from academics and from the general public of Angola. On behalf of all the contributors to this volume, the Editors wish to thank the Ministers, past and present, of Science and Technology, Dr Cândida Teixeira and Dr Maria do Rosário Bragança Sambo, and of Environment, Dr Fátima Jardim and Dr Paula Francisco, for the support given to students of Angola’s biodi- versity. Similarly, the encouragement and logistical support of Prof Liz Matos, Prof Serôdio d’Almeida (Universidade Agostinho Neto), Dr Charles Skinner (De Beers, Angola) and General João Traguedo (Lubango) are gratefully acknowledged. Without their strong support, the scientific results described in this synthesis would not have been possible. Special thanks are due to Martim Melo, who provided his superb copy-editing, proofreading and technical support during the final preparation of the manuscript for submission to the publishers. Similarly, Pedro Tarroso and John Mendelsohn offered their graphic design skills to greatly improve many figures and maps. Photos were also generously offered by the chapter contributors and by Maans Booysen, Merle Huntley, Tassos Leventis, Lars Petersson, Fiona Tweedie and Alexandre Vaz. The financial and logistic support of CIBIO and the UNESCO Chair Life on Land (University of Porto, in association with Twinlab partners in Angola, Namibia, Mozambique, South Africa and Zimbabwe) in this project was fundamental to its success. x Finally, our thanks go to Margaret Deignan, Esther Rentmeester, Malini Arumugam and Maadhuri Kandrakota of Springer (English Edition) and to Jorge Reis-Sa of Arte e Ciência (Portuguese Edition), for their consistent and professional support throughout the project. Acknowledgements xi Contents Part I Introduction: Setting the Scene 1 Angolan Biodiversity: Towards a Modern Synthesis . . . . . . . . . . . . . . 3 Brian J. Huntley and Nuno Ferrand 2 Angola in Outline: Physiography, Climate and Patterns of Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Brian J. Huntley 3 Marine Biodiversity of Angola: Biogeography and Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Stephen P. Kirkman and Kumbi Kilongo Nsingi 4 The Fossil Record of Biodiversity in Angola Through Time: A Paleontological Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 Octávio Mateus, Pedro M. Callapez, Michael J. Polcyn, Anne S. Schulp, António Olímpio Gonçalves, and Louis L. Jacobs Part II Flora, Vegetation and Landscape Change 5 The Flora of Angola: Collectors, Richness and Endemism . . . . . . . . . 79 David J. Goyder and Francisco Maiato P. Gonçalves 6 Vegetation Survey, Classification and Mapping in Angola . . . . . . . . . 97 Rasmus Revermann and Manfred Finckh 7 Suffrutex Dominated Ecosystems in Angola . . . . . . . . . . . . . . . . . . . . . 109 Paulina Zigelski, Amândio Gomes, and Manfred Finckh 8 Landscape Changes in Angola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 John M. Mendelsohn xii Part III Invertebrate Diversity: Environmental Indicators 9 The Dragonflies and Damselflies of Angola: An Updated Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 Jens Kipping, Viola Clausnitzer, Sara R. F. Fernandes Elizalde, and Klaas-Douwe B. Dijkstra 10 The Butterflies and Skippers (Lepidoptera: Papilionoidea) of Angola: An Updated Checklist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167 Luís F. Mendes, A. Bivar-de-Sousa, and Mark C. Williams Part IV Vertebrates: Distribution and Diversity 11 The Freshwater Fishes of Angola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207 Paul H. Skelton 12 The Amphibians of Angola: Early Studies and the Current State of Knowledge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243 Ninda Baptista, Werner Conradie, Pedro Vaz Pinto, and William R. Branch 13 The Reptiles of Angola: History, Diversity, Endemism and Hotspots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283 William R. Branch, Pedro Vaz Pinto, Ninda Baptista, and Werner Conradie 14 The Avifauna of Angola: Richness, Endemism and Rarity . . . . . . . . . 335 W. Richard J. Dean, Martim Melo, and Michael S. L. Mills 15 The Mammals of Angola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357 Pedro Beja, Pedro Vaz Pinto, Luís Veríssimo, Elena Bersacola, Ezequiel Fabiano, Jorge M. Palmeirim, Ara Monadjem, Pedro Monterroso, Magdalena S. Svensson, and Peter John Taylor 16 The Cetaceans (Whales and Dolphins) of Angola . . . . . . . . . . . . . . . . 445 Caroline R. Weir 17 The Giant Sable Antelope: Angola’s National Icon . . . . . . . . . . . . . . . 471 Pedro Vaz Pinto Part V Research and Conservation Opportunities 18 Biodiversity Conservation: History, Protected Areas and Hotspots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495 Brian J. Huntley, Pedro Beja, Pedro Vaz Pinto, Vladimir Russo, Luís Veríssimo, and Miguel Morais Contents xiii 19 Museum and Herbarium Collections for Biodiversity Research in Angola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 513 Rui Figueira and Fernanda Lages 20 Conclusions: Biodiversity Research and Conservation Opportunities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 543 Vladimir Russo, Brian J. Huntley, Fernanda Lages, and Nuno Ferrand Contents xv Contributors Ninda Baptista Instituto Superior de Ciências da Educa ҫ ão da Huíla, Rua Sarmento Rodrigues, Lubango, Angola National Geographic Okavango Wilderness Project, Wild Bird Trust, Parktown, Gauteng, South Africa CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Universidade do Porto, Vairão, Portugal Pedro Beja CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vairão, Portugal CEABN-InBio, Centro de Ecologia Aplicada “Professor Baeta Neves”, Instituto Superior de Agronomia, Universidade de Lisboa, Lisboa, Portugal Elena Bersacola Nocturnal Primate Research Group, Faculty of Humanities and Social Sciences, Oxford Brookes University, Oxford, UK A. Bivar-de-Sousa Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Lisboa, Portugal Sociedade Portuguesa de Entomologia, Lisboa, Portugal William R. Branch (deceased) National Geographic Okavango Wilderness Project, Wild Bird Trust, Parktown, Gauteng, South Africa Department of Zoology, Nelson Mandela University, Port Elizabeth, South Africa Pedro M. Callapez CITEUC; Departamento de Ciências da Terra, Faculdade de Ciências e Tecnologia, Universidade de Coimbra, Coimbra, Portugal Viola Clausnitzer Senckenberg Museum for Natural History, Görlitz, Görlitz, Germany xvi Werner Conradie National Geographic Okavango Wilderness Project, Wild Bird Trust, Hogsback, South Africa School of Natural Resource Management, Nelson Mandela University, George, South Africa Port Elizabeth Museum (Bayworld), Humewood, South Africa W. Richard J. Dean DST-NRF Centre of Excellence at the FitzPatrick Institute, University of Cape Town, Rondebosch, South Africa Klaas-Douwe B. Dijkstra Naturalis Biodiversity Center, Leiden, The Netherlands Sara R. F. Fernandes Elizalde SASSCAL – BID GBIF, Instituto de Investigação Agronómica, Huambo, Angola Ezequiel Fabiano Department of Wildlife Management and Ecotourism, Katima Mulilo Campus, Faculty of Agriculture and Natural Resources, University of Namibia, Katima Mulilo, Namibia Nuno Ferrand CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Campus de Vairão, Universidade do Porto, Vairão, Portugal Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, Porto, Portugal Department of Zoology, Auckland Park, University of Johannesburg, Johannesburg, South Africa Rui Figueira CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vairão, Portugal CEABN-InBio, Centro de Ecologia Aplicada “Professor Baeta Neves”, Instituto Superior de Agronomia, Universidade de Lisboa, Lisboa, Portugal Manfred Finckh Institute for Plant Science and Microbiology, University of Hamburg, Hamburg, Germany Amândio Gomes Institute for Plant Science and Microbiology, University of Hamburg, Hamburg, Germany Faculty of Sciences, Agostinho Neto University, Luanda, Angola António Olímpio Gonçalves Departamento de Geologia, Faculdade de Ciências, Universidade Agostinho Neto, Luanda, Angola Francisco Maiato P. Gonçalves National Geographic Okavango Wilderness Project, Wild Bird Trust, Parktown, Gauteng, South Africa Instituto Superior de Ciências da Educa ҫ ão da Huíla, Lubango, Angola University of Hamburg, Institute for Plant Science and Microbiology, Hamburg, Germany Contributors xvii David J. Goyder Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, UK National Geographic Okavango Wilderness Project, Wild Bird Trust, Parktown, Gauteng, South Africa Brian J. Huntley CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vairão, Portugal Louis L. Jacobs Roy M. Huffington Department of Earth Sciences, Southern Methodist University, Dallas, TX, USA Jens Kipping BioCart Ökologische Gutachten, Taucha/Leipzig, Germany Stephen P. Kirkman Department of Environmental Affairs, Oceans and Coasts Research, Cape Town, South Africa Fernanda Lages ISCED – Instituto Superior de Ciências da Educação da Huíla, Lubango, Angola Octávio Mateus GeoBioTec, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, Lisbon, Portugal Museu da Lourinhã, Rua João Luis de Moura, Lourinhã, Portugal Martim Melo DST-NRF Centre of Excellence at the FitzPatrick Institute, University of Cape Town, Rondebosch, South Africa CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Universidade do Porto, Vairão, Portugal Instituto Superior de Ciências da Educa ҫ ão de Huíla, Lubango, Angola John M. Mendelsohn RAISON (Research & Information Services of Namibia), Windhoek, Namibia Luís F. Mendes Museu Nacional de História Natural e da Ciência, Universidade de Lisboa, Lisboa, Portugal CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Vairão, Portugal Michael S. L. Mills Instituto Superior de Ciências da Educa ҫ ão de Huíla, Lubango, Angola A. P. Leventis Ornithological Research Institute, University of Jos, Jos, Plateau State, Nigeria Ara Monadjem Department of Biological Sciences, University of Swaziland, Kwaluseni, Swaziland Mammal Research Institute, Department of Zoology and Entomology, University of Pretoria, Pretoria, South Africa Contributors xviii Pedro Monterroso CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vairão, Portugal Miguel Morais Faculdade de Ciências, Universidade Agostinho Neto, Luanda, Angola Kumbi Kilongo Nsingi Benguela Current Convention, Swakopmund, Namibia Jorge M. Palmeirim Departamento de Biologia Animal, Faculdade de Ciências, cE3c – Centre for Ecology, Evolution and Environmental Changes, Universidade de Lisboa, Lisboa, Portugal Pedro Vaz Pinto Fundação Kissama, Luanda, Angola CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus de Vairão, Vairão, Portugal Michael J. Polcyn Roy M. Huffington Department of Earth Sciences, Southern Methodist University, Dallas, TX, USA Rasmus Revermann Institute for Plant Science and Microbiology, University of Hamburg, Hamburg, Germany Vladimir Russo Fundação Kissama, Luanda, Angola Anne S. Schulp Naturalis Biodiversity Center, Leiden, The Netherlands Faculty of Earth and Life Sciences, VU University Amsterdam, Amsterdam, The Netherlands Paul H. Skelton South African Institute for Aquatic Biodiversity (SAIAB), Grahamstown, South Africa Wild Bird Trust, National Geographic Okavango Wilderness Project, Hogsback, South Africa Magdalena S. Svensson Nocturnal Primate Research Group, Faculty of Humanities and Social Sciences, Oxford Brookes University, Oxford, UK Peter John Taylor School of Mathematical & Natural Sciences, University of Venda, Thohoyandou, South Africa Luís Veríssimo Fundação Kissama, Luanda, Angola Caroline R. Weir Ketos Ecology, Kingsbridge, Devon, UK Mark C. Williams Pretoria University, Pretoria, South Africa Paulina Zigelski Institute for Plant Science and Microbiology, University of Hamburg, Hamburg, Germany Contributors Part I Introduction: Setting the Scene 3 © The Author(s) 2019 B. J. Huntley et al. (eds.), Biodiversity of Angola , https://doi.org/10.1007/978-3-030-03083-4_1 Chapter 1 Angolan Biodiversity: Towards a Modern Synthesis Brian J. Huntley and Nuno Ferrand Abstract Angola possesses an unusually rich diversity of ecosystems and species, but this natural wealth is poorly documented when compared with other countries in the region. Both colonial history and extended wars challenged progress in biodiver- sity research and conservation, but since peace was achieved in 2002 a rapidly increasing level of collaboration between Angolan and visiting scientists and institu- tions has seen a blossoming of biodiversity research. The absence of comprehensive reviews and syntheses of existing knowledge, often published in extinct journals and inaccessible official reports, necessitates a modern synthesis. This volume brings together the existing body of scientific results from studies on Angola’s landscapes, ecosystems, flora and fauna, and presents an outline of opportunities for biodiversity discovery, understanding and conservation as well as collaborative research. Keywords Africa · Biomes · Collaborative research · Conservation · Ecoregions Background and Context Angola is a country of unusually rich physiographic, climatic and biological diver- sity. It occupies only 4% of the terrestrial area of Africa, yet it possesses the highest diversity of biomes and is second only to mega-diverse South Africa in terms of the number of ecoregions found within its borders. However, scientific literature on its B. J. Huntley ( * ) CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vairão, Portugal e-mail: brianjhuntley@gmail.com N. Ferrand CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Campus de Vairão, Universidade do Porto, Vairão, Portugal Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, Porto, Portugal Department of Zoology, Auckland Park, University of Johannesburg, Johannesburg, South Africa e-mail: nferrand@cibio.up.pt 4 biodiversity is extremely limited when compared with most African countries. Much of that which has been published is difficult to access or out of print. This volume seeks to redress this situation. Here we present a review of what is known about Angola’s biodiversity. Much of the existing literature dates from the nineteenth and early to mid-twentieth centu- ries. Following independence in 1975, field studies were curtailed by the instabili- ties of an extended civil war. It was not until after the peace settlement of 2002 that a new wave of research has been possible. Initial attempts to establish collaborative field expeditions were frustrated by visa and permit restrictions, but these chal- lenges were gradually overcome and by the 2010s a vibrant programme of joint projects has evolved. Today many foreign specialists work in partnership with Angolan researchers and institutions, producing a new flow of scientific results of which many are presented in this volume. For any comprehensive synthesis, both temporal depth and spatial breadth is necessary. An historical perspective is presented in each chapter. Angolan indige- nous knowledge has contributed to the insights and materials that have informed visiting researchers from the eighteenth century to the present day. The pioneering studies and exhaustive botanical collections of the Austrian botanist, Friedrich Welwitsch (1806–1872), the zoological collections of the indefatigable Portuguese naturalist José Anchieta (1832–1897) and the Swiss botanist John Gossweiler (1873–1952) set benchmarks for later work (Swinscow 1972; de Andrade 1985). Each succeeding student of Angola has added to the description of its biological diversity. While botanists such as Romero Monteiro (1970) and zoologists such as Crawford-Cabral (1983) have summarised available biogeographic information within a national context, no comprehensive synthesis of studies on Angola’s fauna, flora and ecosystems has yet been undertaken. The need for an integrated account has become evident in the past decade, as increasing numbers of expeditions and collaborative projects have evolved as part of the country’s ‘peace dividend’. Approach and Purpose of This Synthesis A modern synthesis is not easily achieved. Much of the early literature on Angola’s biodiversity resides in publications and reports that are difficult to source. This review attempts to reference these important but sometimes elusive accounts, in order to provide students with access to what information is available. While focus- ing on papers in peer-reviewed journals, some topics need to draw on unpublished reports filed in government departments. It also seeks to bring together the findings of recent, post-independence studies, many of which are still in progress or in press. It is intended to serve the new generation of Angolan students by providing a com- prehensive but focused synopsis of what is known on the biomes, landscapes, flora and fauna of Angola. It should also bring Angola to the attention of researchers across Africa and beyond, revealing the great diversity of life, and the multiple questions on the structure and functioning of Angola’s biodiversity that await explo- ration, examination and explanation. B. J. Huntley and N. Ferrand 5 In structuring this book, this introduction leads through synopses on the coun- try’s terrestrial and marine biogeography, paleontological record, recent landscape evolution and land transformation, to chapters on its flora and vegetation. The main body of this volume is devoted to accounts of its fauna – selected invertebrate groups that have promise as indicators of environmental stress, and all vertebrate groups. In each treatment, the need for increased conservation measures for threatened taxa and habitats is a recurrent theme, while research opportunities are highlighted. While general inventories and checklists are progressing well, the state of ecologi- cal knowledge remains rudimentary. Topics as fundamental as ecological processes such as the flows of energy, water and nutrients; the ecological impacts of phenom- ena such as fire, invasive species, herbivory, droughts and frosts; community struc- ture, plant-animal interactions and the impacts of land-transformation and of climate change are yet to be researched in Angola. This volume’s content is limited by the availability of information. It is therefore opportunistic, covering those taxonomic groups and those features and processes for which a critical mass of information is available. The focus is primarily on the terrestrial ecosystems and biota of Angola, but the importance of the marine environment is described in accounts on marine biodiversity and ocean dynamics, and on the richness of the whale, dolphin and marine turtle faunas of Angolan waters. In comparison with similar reviews for other African countries with long and strong traditions of research into their biodiversity and ecology, and for which com- prehensive syntheses of the state of knowledge are available (e.g. Namibia: Barnard 1998; Southern Africa: Davis 1964; Werger and van Bruggen 1978; Huntley 1989; Tanzania: Sinclair 2012), this account reveals both the strengths and weaknesses of the research agenda of the colonial era, and the challenges of the recent past. While institutions such as the Instituto de Investigação Científica de Angola and the Instituto de Investigação Agronómica de Angola undertook very important studies on many taxa, and on vegetation, soils and agronomy, and the Museu do Dundo amassed and distributed a vast series of collections of the animal species of the Lundas, the coverage of disciplines and of the remote regions of Angola was weak. Biodiversity Surveys: Historical Synopsis The history of scientific exploration and biological collection in Angola is relatively modest. Whereas South Africa, by 1975, had over three million herbarium speci- mens collected by some 2500 botanists since the late eighteenth century (Gunn and Codd 1981), Angola had less than 300,000 specimens collected by just 300 bota- nists during the same period (Figueiredo and Smith 2008). Despite the relatively limited coverage of Angolan collections, the great botanist Francisco Mendonça was occasioned to state in his preface to Gossweiler and Mendonça (1939): We are happily able to confirm that the flora of Angola is the best known in tropical Africa, due to the attention given by the state towards the botanical exploration of the colony, and the great interest and zeal of scientists in its study. 1 Angolan Biodiversity: Towards a Modern Synthesis