Brian J. Huntley · Vladimir Russo Fernanda Lages · Nuno Ferrand Editors Biodiversity of Angola 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 United Nations UNESCO Chair on Life on Land Educational, Scientific and University of Porto Cultural Organization Porto, Portugal Editors Brian J. Huntley Vladimir Russo CIBIO-InBIO, Centro de Investigação em Fundação Kissama Biodiversidade e Recursos Genéticos Luanda, Angola Universidade do Porto Vairão, Portugal Nuno Ferrand CIBIO-InBIO, Centro de Investigação em Fernanda Lages Biodiversidade e Recursos Genéticos ISCED – Instituto Superior de Ciências da Laboratório Associado, Campus de Vairão Educação da Huíla Universidade do Porto Lubango, Angola Vairão, Portugal Departamento de Biologia, Faculdade de Ciências Universidade do Porto Porto, Portugal Department of Zoology, Auckland Park University of Johannesburg Johannesburg, South Africa 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. This book is an open access publication 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. The images or other third party material in this book are included in the book’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the book’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland 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 Foreword His Excellency the President of the Republic of Angola, João Manuel Gonçalves Lourenço 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 vii viii Foreword 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 João Manuel Gonçalves Lourenço Luanda, Angola 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. ix x Acknowledgements 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. 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 xi