THE DIVERSITY of LIFE in AFRICAN FRESHWATERS: UNDERWATER, UNDER THREAT an Analysis of the Status and Distribution of Freshwater Species Throughout Mainland Africa

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THE DIVERSITY OF LIFE IN AFRICAN FRESHWATERS: UNDERWATER, UNDER THREAT An analysis of the status and distribution of freshwater species throughout mainland Africa

Edited by: William Darwall, Kevin Smith, David Allen, Robert Holland, Ian Harrison and Emma Brooks

The IUCN Red List of Threatened Species™ – Regional Assessment

THE DIVERSITY OF LIFE IN AFRICAN FRESHWATERS: UNDERWATER, UNDER THREAT An analysis of the status and distribution of freshwater species throughout mainland Africa

Edited by: William Darwall, Kevin Smith, David Allen, Robert Holland, Ian Harrison and Emma Brooks

Project Partners:

Financial support provided by:

and:

In kind funding was provided by: The designation of geographical entities in this book, and the presentation of material, do not imply the expression of any opinion whatsoever on the part of IUCN or other participating organizations, concerning the legal status of any country, territory, or area, or of its authorities, or concerning the delimitation of its frontiers or boundaries.

This project has been carried out with financial support from the European Union under grant Contract: EuropeAid/ENV/2004-81917. The views expressed herein can in no way be taken to reflect the official opinion of the European Union.

The views expressed in this publication do not necessarily reﬂect those of IUCN or other participating organizations.

Published by: IUCN, Gland, Switzerland and Cambridge, UK Copyright: © 2011 International Union for Conservation of Nature and Natural Resources Reproduction of this publication for educational or other non-commercial purposes is authorized without prior written permission from the copyright holder provided the source is fully acknowledged. Reproduction of this publication for resale or other commercial purposes is prohibited without prior written permission of the copyright holder.

Red List logo: © 2008 Citation: Darwall, W.R.T., Smith, K.G., Allen, D.J., Holland, R.A, Harrison, I.J., and Brooks, E.G.E. (eds.). 2011. The Diversity of Life in African Freshwaters: Under Water, Under Threat. An analysis of the status and distribution of freshwater species throughout mainland Africa. Cambridge, United Kingdom and Gland, Switzerland: IUCN. xiii+347pp+4pp cover. ISBN: 978-2-8317-1345-8 Cover design: Cambridge Publishers Cover photos: Front: Tanganyika cichlid (Xenotilapia melanogenys) a Data Deﬁcient species endemic to Lake Tanganyika © Angel M Fitor/Photolibrary Ltd. Back: A girl collects water supplied by Bale Mountains National Park in south-central Ethiopia © Robin Moore All photographs used in this publication remain the property of the original copyright holder (see individual captions for details). Photographs should not be reproduced or used in other contexts without written permission from the copyright holder.

Layout by: Cambridge Publishers Produced by: Cambridge Publishers Available from: IUCN (International Union for Conservation of Nature), Publications Services, 28 Rue Mauverney, 1196 Gland, Switzerland, Tel: + 41 22 999 0000, Fax: + 41 22 999 0020, Email: [email protected], www.iucn.org/publications A catalogue of IUCN publications is also available. INTRODUCTION | CONTENTS iii 4 6 8 ix xi xii 34 34 35 35 36 37 38 39 41 44 44 49 49 49 55 60 63 65 13 13 16 18 22 23 24 25 25 26 26 28 34 34 ......

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...... Habitat transformation Pollution Invasive alien species Impact of ﬁ sheries Impact of ﬁ Climate change

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A short introduction to the ichthyofaunal regions A short introduction General overview ecosystems to African freshwater Threats Species diversity biodiversity to freshwater Major threats 1.2.2.1 1.2.2.2 1.2.2.4 The African Green Revolution and wetlands The African Green ......

Odonates Crabs Aquatic plants sh species All ﬁ species Threatened Restricted Range species cient species Data Deﬁ Extirpated species Ecosystem functions Ecosystem values Fishes Molluscs

Global status of freshwater biodiversity Global status of freshwater 2.1.3 2.1.4 2.1.5 3.1.1 3.3.1 3.3.2 3.3.3 3.3.4 3.3.5 1.2.1 1.2.2 1.2.2.3 1.2.2.5 1.2.3 ecosystems in Africa Functions and values of freshwater 1.3.1 2.1.1 2.1.2 1.1.1 Overview of the ichthyofauna of Africa Selection of priority taxa Regional approach Regional Data collation and quality control mapping Species status Assessment of species threatened Nomenclature Conservation status Patterns of overall species richness Situation analysis for Africa 1.1 Acknowledgements Chapter 1 - Background 2 Contents Introduction Chapter 3 - Fish 3.1 42 2.1 Chapter 2 - Assessment methodology Chapter 2 - Assessment methodology 32 2.2 2.3 2.4 2.5 2.6 3.2 3.3 1.3 1.3.2 Foreword Executive Summary 1.1.2 1.2 iv INTRODUCTION | CONTENTS Species inthespotlight Species inthespotlight 4.5 4.4 4.3 4.2 4.1 Species inthespotlight Species inthespotlight Species inthespotlight Species inthespotlight Species inthespotlight 3.6 3.5 itr,dvriy itiuin n osrain 126 5.1 history, diversity, distribution, andconservation Chapter 5- 92 Introduction diversity, distribution,andconservation Chapter 4-Freshwater molluscsofAfrica: 3.4

Species inthespotlight Species inthespotlight Recommendedconservationmeasures Main threats Overviewoffreshwater molluscandiversityinAfrica Conservationrecommendations Research actionsrequired Majorthreats tospecies

Introduction

Chapter 4-Speciesinthespotlight Chapter 3-Speciesinthespotlight Conservation statusanddistributionoffreshwater molluscs 5.1.2 5.1.1 4.5.1 Understanding theimpactofmolluscsonhumanhealthandlivelihoods 4.3.8 4.3.7 4.3.6 4.3.5 4.3.4 4.3.3 4.3.2 4.3.1 4.2.5 4.2.4 4.2.3 4.2.2 4.2.1 4.1.6 4.1.5 4.1.4 4.1.3 4.1.1 4.1.2 Knowledge ofAfricanOdonata Odonatainbiogeography andconservation Recommendedresearch actions Invasivespecies–fi sh farming Sedimentation Water pollution ofthreat Regionalpatterns Extinctionpatterns DataDefi cient species Threatened species Molluscandiversityincraterlakes Molluscandiversityinartesianbasins Molluscandiversityinriverbasins Molluscandiversityinancientlakes 4.2.1.1 Climate change Invasive species–molluscsandtheirimpactonindigenous Invasive species–waterhyacinthandotherplants Dams Undescribed species Restricted rangespecies Molluscan diversityinmontanelakes Molluscan diversityinsalinelagoons,saltmarshesandmangroves ...... Dragonfl ...... Changessincethe1996assessment ...... –Amolluscwithabitofmuscle –Canthelastpopulationsof giantAfricanfreshwater pearlmusselbesaved? –Cauldrons forfi Africa’ssh biodiversity:western craterlakes –TheTwee Riverredfi n –aCriticallyEndangered minnowfrom SouthAfrica –Auniquespeciesfl ock inLakeTana –theLabeobarbuscomplex –Forest remnantsAfrica–vanishingislands ofsylvanfi inwestern shes Africa–afriendandfoe ineastern –Tilapia – – ...... Freshwater molluscsare important–especially totheAfricanopenbillstork The Congoblindbarb:MbanzaNgungu’s albinocavefi sh ...... ies anddamselfl ......

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...... ies ofAfrica(Odonata): ......

...... 128 128 128 124 123 120 119 118 117 117 114 114 114 114 114 113 112 112 111 110 109 106 106 105 104 104 103 102 102 100 97 96 94 87 85 82 79 76 74 71 70 66 INTRODUCTION | CONTENTS v 166 166 167 168 168 169 169 170 170 170 170 171 172 172 173 174 175 176 177 128 129 132 132 132 132 133 133 133 138 141 143 143 145 145 147 148 149 153 153 154 156 156 158 159 159 160 162 162 164 164 166 ......

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...... A guardian of Kenya's watershed: Maathai's longleg of Kenya's A guardian ...... – – The globe skimmer: on a journey the world around y in a fragile realm – The Pemba featherleg: a fragile damselfl with an ancient lineage – A wonderful water relic ...... Taxonomy and identifi cation and identifi Taxonomy Red List assessment of African dragonfl ies Red List assessment of African dragonfl Dams Understanding change Distribution and ecology and Distribution Patterns of diversity cation Impoverishment and diversifi Relict fauna Modern fauna Upper Guinea Lower Guinea Congo Basin Overview South Africa Angola Eastern Mountains and Coast Indian Ocean Atlantic Ocean ...... species Distribution of threatened Habitat degradation and deforestation Mining sh Alien plants and fi extraction and pollution Water Field survey Ecological research Information disclosure Richness Relevance and context Overview Ethiopia Albertine Rift ...... Zambezia ...... Major threats Diversity fauna Afrotropical fauna Afro-montane Insular fauna 5.1.2.1 5.1.2.1 5.4.1.1 5.4.1.2 5.4.2.1 5.4.2.2 5.4.2.3 5.4.2.4 5.4.2.5 5.4.3.1 5.4.3.2 5.4.3.3 5.4.3.4 5.1.2.2 5.1.2.2 5.2.1.1 5.2.1.2 5.2.2.1 5.2.2.2 5.2.2.3 5.2.2.4 5.3.2.1 5.3.2.2 5.3.2.3 5.3.2.4 5.3.3.1 5.3.3.2 5.3.3.3 5.3.3.4 5.3.3.5 5.3.4.1 5.3.5.1 5.3.5.2 Research actions Research Conservation actions History Guineo-Congolian fauna Complex faunas fauna Palaearctic Status 5.4.2 5.4.3 Diversity and history Diversity and 5.2.1 5.2.2 endemism and threat Regional diversity, 5.3.1 5.3.2 5.3.3 5.3.4 5.3.4.2 5.3.5 5.3.6 Conservation 5.4.1 Chapter 5 - Species in the spotlight

Species in the spotlight

5.4.4 Species in the spotlight Species in the spotlight Species in the spotlight 5.2 5.3 5.4 vi INTRODUCTION | CONTENTS 7.8 7.7 7.6 7.5 7.4 7.3.2 7.3 7.2 Species inthespotlight 6.6 6.5 6.4 6.3 6.2 hpe qai lnso fia iest,dsrbto n osrain 200 Chapter 7-AquaticplantsofAfrica:diversity, distributionandconservation 6.1 178 Chapter 6-Freshwater crabsofAfrica:diversity, distribution,andconservation 7.1 Species inthespotlight Keymessages Conservationrecommendations ofspeciesrichness Patterns Extirpatedspecies Conservationrecommendations Majorthreats ofspeciesrichness Patterns Conservationstatus Background Majorthreats to aquatic plants Research actionsrequired

Chapter 6-Speciesinthespotlight Conservation 7.7.6 7.7.5 7.7.4 7.7.3 7.7.2 7.7.1 7.6.7 7.6.6 7.6.5 7.6.4 7.6.3 7.6.2 7.6.1 7.5.4 7.5.3 7.5.2 7.5.1 7.3.1 Overview ofthefreshwater fl ora ofmainlandcontinentalAfrica 6.4.4 6.4.3 6.4.2 6.4.1 6.3.4 6.3.3 6.3.2 6.3.1 6.1.1 Overview oftheAfricanfreshwater crabfauna Endemicspecies Furtherresearch Speciesorsitebasedactions Alienspeciesremoval Planningandpolicy Climatechange Invasivespecies Water use:extractionandtransfers DataDefi cient species Threatened species Continentalendemicspecies Allassessedspecies Allspecies Taxonomic issues Pollution Habitatdestruction DataDefi cient species Restrictedrangespecies Biogeographicpatterns Restoration Catchment-scale actions Water pollution Habitat lossduetoinfrastructure development Habitat lossduetoagriculture General threats Natural predators andcompetitionwithintroduced species Threatened species All freshwater crabspecies:interpretation ofdistributionpatterns ...... tatus s ......

...... –TheEastAfricantree holecrab–spectacularadaptation – ......

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...... studies inevolutionandextinction A uniquespeciesﬂ ock offreshwater crabsinLakeTanganyika: amodelfor ......

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...... 223 223 222 221 221 221 221 221 221 221 220 219 218 216 216 215 214 211 209 209 209 209 208 205 205 204 202 198 196 195 193 193 192 192 191 191 190 188 187 186 184 183 182 180 INTRODUCTION | CONTENTS vii 272 272 272 274 274 275 275 277 277 278 278 278 282 282 283 284 284 286 286 224 225 226 227 230 230 230 235 244 247 247 247 249 249 251 258 258 258 262 264 264 265 265 267 267 268 ......

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...... A small wonder of nature saved from extinction saved from of nature wonder A small Water hyacinth, a threat to the freshwater biodiversity of western Africa to the freshwater hyacinth, a threat Water ...... – – Sailing ﬂ owers – Sailing ﬂ creator – The habitat – ......

...... When are site-scale actions or planning appropriate? site-scale When are Prioritisation of site-scale conservation actions Recent developments General issues of connectivity connectivity and lateral Threats ...... Adapting terrestrial approaches to freshwater ecosystems to freshwater approaches Adapting terrestrial Case studies of site-scale conservation intervention in southern Africa Planning and implementation spatial scales as units for conservation planning ecoregions Freshwater Patterns of richness and main threats species to African freshwater Distribution of threats species in Africa freshwater Biological invasions affecting taxonomic groups Congruence between ecosystems for freshwater areas Protected species for freshwater Key Biodiversity Areas Extinction – AZE sites Alliance for Zero species species and threatened cient Regions of Data Deﬁ of new species cient species with records Comparing distributions of Data Deﬁ (RAPs) Rapid Assessment Programs the past through the future conservation: seeing Climate change and African freshwater African waters: special qualities and challenges 100,000 years dynamism: the past A short history of African eco-hydrological climate change in Africa Signs of anthropogenic and responses trends Projecting change to freshwater Implications for facilitating healthy responses 9.1.3.2 9.1.3.3 9.2.1.1 9.2.1.2 Background ecosystems Application to freshwater Catchment versus site scale approaches Conclusions Use of biodiversity surrogates Data requirements

9.1.1 9.1.2 9.1.3 9.2.1 9.2.2 9.2.3 9.3.1 9.3.2 9.3.3 8.2.1 8.2.2 8.2.3 8.3.1 8.3.2 8.3.3 8.3.4 8.5.1 8.5.2 8.5.3 8.6.1 8.6.2 8.6.3 8.6.5 8.6.6 Chapter 7 - Species in the spotlight in the 7 - Species Chapter

Introduction Conservation action Conservation Systematic conservation planning Site scale conservation Catchment scale conservation Conclusions Africa biodiversity across Freshwater species Livelihoods dependence on freshwater for further work Areas of African freshwaters The future Species in the spotlight in the Species Chapter 9 - Conservation of freshwater ecosystems Chapter 9 - Conservation of freshwater 9.1 270 8.1 Chapter 8 - Synthesis for all taxa Chapter 8 - Synthesis 228 9.1.3.1 9.2 9.3 9.3.4 9.4 Species in the spotlight Species in the spotlight Species in the spotlight Species in 8.2 8.3 8.4 8.5 8.6 8.6.4 viii INTRODUCTION | CONTENTS Chapter 10References Chapter 9References Chapter 8References Chapter 7References Chapter 6References Chapter 5References Chapter 4References Chapter 3References Chapter 2References 10.2 An exampleofaspeciesassessmentsummaryanddistributionmap Appendix Chapter 1References References 288 10.1 Chapter 10-Developmentandpolicyplansforthefuture Freshwater biodiversitymanagementandconservation, policy, andtheprivatesector Wetland developmentplansforAfrica 10.2.3 10.2.2 10.2.1 10.1.5 10.1.4 10.1.3 10.1.2 10.1.1 Policyandtheprivatesector–conclusion Roleoftheprivatesector Thepolicyprocess Threat mitigationandmanagementplansthatintegrate freshwater biodiversity Impactofwetlanddevelopmentoninlandwaters DegradationandlossofAfrica’s wetlands Wetland resources anduses Development across Africa

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...... 346 306 346 343 339 335 333 330 325 322 313 312 306 305 302 299 299 297 297 292 291 290 290 INTRODUCTION Acknowledgements |

Donors his valuable and thorough review of the chapter on plants, and IUCN would like to thank all the donors that have made this to C.D.K. Cook and H.J. Beentje for their useful comments. ACKNOWLEDGEMENTS assessment possible. The project’s major donor was the Tom Brooks, Will Turner, and Daniel Brito are thanked for European Commission (EC), under grant number EuropeAid/ providing access to relevant unpublished materials. ENV/2004-81917. Financial support was also provided by the BioFresh project (funded by the European Union under the 7th Report photos Framework programme, Contract No. 226874); Conservation We would also like to thanks those who have contributed International (CI); Darwin Initiative of the UK Department for photos at no charge, including Conservation International Environment, Food and Rural Affairs (Defra); IUCN National Marketing and Communications, and the numerous Committee of the Netherlands; IUCN Water and Nature photographers who have provided access to their images Initiative (WANI); MAVA Foundation pour la nature (MAVA); through Creative Commons. Ministrie von Buitenlandse Zaken/Dutch Ministry of Foreign Affairs (DGIS); Spanish Agency for International Cooperation GIS expertise Development (AECID); and United Nations Environment The species distribution mapping and analysis would not Programme World Conservation Monitoring Centre (UNEP- have been possible without the GIS expertise provided by: 1) WCMC). In-kind support was provided by the Junta de IUCN; Vineet Katariya; Adrian Hughes; Yichuan Shi; Jemma Andalucia; Ministerio de Medio Ambiente y Medio Rural y Able; and Sussanah O’Hanlon; and 2) UNEP-WCMC; Simon Marino; Okavango Research Institute (ORI), formerly the Blyth; Igor Lysenko and Corinna Ravilious. We also thank Harry Oppenheimer Okavango Research Centre (HOORC); the many volunteers who helped with mapping and editing South African National Biodiversity Institute (SANBI); South species distributions: Amy Milam; Arabie Jaloway; Ian Denty; African Institute for Aquatic Biodiversity (SAIAB); Tour du Jennifer Taylor; Jo Fisher; Joe Wood; John Lynch; Kasia Valat; and Wetlands International (WI). Brooks; Kayshinee Ramchun; Pramod Lamsal; and Sarah Deacon. Project partners IUCN Species Programme, through its Freshwater Biodiversity IUCN Species Programme Unit, led this project in close collaboration with the following We thank the following members of the IUCN Species project partners, without which the project would not Programme who have provided support and training at have been possible: the South African Institute for Aquatic workshops and who also manage the IUCN Red List Biodiversity (SAIAB); the IUCN Centre for Mediterranean database and website through which the project assessment Cooperation; Wetlands International; the IUCN Regional data are made available: Anabelle Cuttelod; Caroline Pollock; Ofﬁ ce for Southern and Eastern Africa (IUCN ESARO); ITC Craig Hilton-Taylor; Helen Temple; Julie Grifﬁ n; Leah Collett; (Faculty of Geo-Information Science and Earth Observation, Maiko Lutz; Melanie Bilz; and Rebecca Miller. We also University of Twente); and the United Nations Environment thank Jane Smart, as Head of the Species Programme, Programme World Conservation Monitoring Centre (UNEP- for her support and advice throughout this project. James WCMC). Ragle, who manages the Species Information Services (the database behind the IUCN Red List), is also acknowledged. Steering Committee Thanks are also due to the following members of the IUCN The members of Project Steering Committee are Communications teams who have provided their expertise acknowledged for providing sound knowledge and guidance on press releases and publications: Lynne Labanne; Kathryn throughout the life of the project. Annabelle Cuttelod, Pintus; Chadi Abi Faraj; and Sonsoles San Roman. We would Christoph Zöckler, Edward Westinga, Eliot Taylor, Fabio also like to thank Anna McIvor, Laurel Bennett, Nieves Garcia, Corsi, Geoffrey Howard, Hein van Gils, Ibrahima Thiam, Jean- and Thomas Lowe, who are no longer with IUCN Freshwater Christophe Vié, Jessica Jones, Laurent Ntahuga, Madiodio Biodiversity Unit but have contributed in many important Niasse, Mamadou Niane, Mame Dagou Diop, Matt Walpole, ways to the success of this project. Finally, we would like Neville Ash, Nkobi Moleele, Patrick Van Laake, Paul Skelton, to thank Jean-Christophe Vié for his valued support and Rami Salman, and Ward Hagemeijer. oversight throughout the project.

Report contributors Finances The editors thank the authors and contributors to this report, The fi nances of the project have been diligently managed and those others who have commented on the various stages by Maureen Martindell, Verity Saunders, Hugo Ruiz, Josette of the manuscript. Very special thanks to J.R. Timberlake for Lehmann and Line Hempel, who all deserve great credit, ix as do the fi nance teams of the project partners, including H. Brett, E. Brooks, R. Brummett, P. Budot, J. Burrows, J. Ronald Hasselerharm (ITC); Damas Mugashe, Thaddeus Cambray, B. Celestin, A.B. Cham, M. Cheek, A. Chilala, Kamundi and Edward Mudida (IUCN ESARO); Esther S. Chimatiro, F. Cholo, V. Clausnitzer, F.-L. Clotilde-Ba, W. Adhiambo Abonyo (IUCN Global Invasive Species Initiative); Coetzer, B. Collen, S. Conde, T. Contreras-MacBeath, A.J. Mohamed Karmass (IUCN Mediterranean Offi ce); Jane Crivelli, N. Cumberlidge, C. Cupido, B. Curtis, A. Cuttelod, Stockwell (SAIAB); Alex Gee (UNEP-WCMC); and Guenoune L. Da Costa, L. Dacunha, M. Dakki, F. Daniels, S. Daniels, Fatou Mbengue (Wetlands International Africa). A. Daoud-Bouattour, F.S. Darboe, W.R.T. Darwall, J. Day, L. De Vos, M.P. de Zeeuw, R. Dessouassi, C.H. Diagana, M.S.

ACKNOWLEDGEMENTS European Commission support Diallo, M. Diedhiou, E. Dieme-Amting, K.-D.B. Dijkstra, A.L.

| We would also like to thank Joan Cummins, who provided us Diop, F.N. Diop, M.D. Diop, M.S. Diop, A. Diouf, A.P. Dold, with guidance and support from the European Commission. R. Dow, L. Dreyer, I. Ebrahim, A.R. El Gamal, H.S. Elgrari , S. Elongo, J. Engelbrecht, M. Entsua-Mensah, H. Ferchichi, M.J. DVD layout Fernández, S.A.F. Ferreira, L. Fish, M.R.A. Fishar, D. Flanagan, The enclosed data DVD was produced using a programme W. Foden, S. Fofana, A.R. Gamal, Z. Gammar-Ghrabi, N. kindly written by Nicola Terry. Garcia, P. Gateretse, A. Getahun, M. Ghamizi, J. Ghogue, Z. Ghrabi, N. Gichuki, D. Graff, J. Griffi n, P. Grillas, M.B. Gueye, INTRODUCTION Logistics F. Habineza, O.E. Hadi, W. Hagemeijer, E. Hakizimana, M. We have, throughout the project, also been provided logistical Hanssens, B. Hassiba, C. Hay, N. Helme, C. Hilton-Taylor, support from non-project partners, and we would like to give G. Howard, H. Howege, L. Hugot, M. Hussein, S.N.B. Ifuta, our thanks to them: Samy Zalat, Randal Brummett and staff D. Impson, O.O. Itoua, G. Jacobseni, G. Jakubowsky, M.S. (WorldFish Centre), and The Research Centre for Biodiversity Jallow, H. James, I.B.H. Jilani, D. Johnson, A. Jørgensen, and Genetic Resources of Porto University (CIBIO-UP). B. Joseph, M. Jovic, A. Kabuto, A. Kagabo, D. Kahwa, V. Kalkman, D.A. Kamundi, A. Kane, R. Kane, C. Kapasa, M.H. IUCN Specialist Groups Kara, J. Kasembe, E. Kaunda, J. Kazembe, D. Khawa, N. The project has worked very closely with the IUCN Species Khumalo, J. Kipping, J. Kisakye, Z. Kiza, M. Kraiem, T.K. Survival Commission Specialist Group network who often Kristensen, J. Kruger, F. Krupp, R. Kyambadde, P. Laleye, provided species expertise. We would like to thank the IUCN/ C. Lange, S. Limam-Ben Saad, M. Lock, H. Lockwood, T. Wetlands International Freshwater Fish Specialist Group; Lowe, W.R.Q. Luke, P. Mafabi, Z. Magombo, A. Mahamane, Mollusc Specialist Group; Dragonfl y Specialist Group, and A. Mailosa, P. Makocho, V. Mamonekene, M.M. Manga, J. the Freshwater Crab and Crayfi sh Specialist Group. Manning, P.A. Manyama, S.A.E. Marijnissen, E. Marinus, B. Marshall, P.F. Matlamela, P.O. Mbeke, A. Mcivor, F. Médail, Additional regional acknowledgments A. Melembe, E. Michel, A. Mitra, T. Moelants, G. Mokhtar, A series of regional reports have also been published for which N. Mollel, S. Mondal, K.A. Monney, R.J.C. Monsembula, many additional people and organisations provided support; F.De Moor, A. Moyo, J. Msibi, Mubbala, A. Muhweezi, K. please see these reports for their acknowledgements. In Mukungilwa, S. Müller, J. Musoni, M. Mutekanga, B. Mwangi, particular, we thank the participation and contributions K. Naidoo, G. Natakimazi, A. Ndiaye, D.S. Ndiaye, T. Ndiaye, by many representatives of national scientifi c institutions G. Ndiritu, B. Ndodet, L. Nel, B. Ngatunga, C. Ngereza, and government agencies who attended workshops and B.K. Ngoy, F. Nicayennzi, N. Nje, K.A. Noby, M.V. Nshombo, supported the project. G. Ntakimazi, J. Ntukamazina, P. Ntumigomba, M. Nyaligu, T. Nzabi, L. Nzeyimana, K. Oberlander, G. O’Brien, E.A. Assessors, evaluators and workshop Odhiambo, S.S. Ogbogu, B. Olaosebikan, J.M. Onana, M. participants Ouali, L. Ouédraogo, D. Oyugi, D. Paugy, D. Petrovic, D. Pillay, Last, but defi nitely not least, the heart of this project is the C. Pollock, L. Potter, V. Pouomogne, V. Prié, D. Raimondo, M. information on African freshwater biodiversity that has been Ram, G.M. Reid, L. Rhazi, M. Rhazi, R. Riddick, E. Riservato, provided by many enthusiastic and committed experts from J.P. Roux, E. Rufuguta, J. Sa, S.B. Saad, S.L. Saad, A.M. across Africa and the wider world. Without these individuals Sajer, F. Salgado, B. Sambou, B. Samraoui, M. Samways, E. this work would not be possible, and they deserve special Sattout, W. Schneider, E. Schraml, K. Schütte, C.R. Scott- acknowledgement. Shaw, A. Seck, M. Seddon, S. Senol, C. Sheifer, A. Shmida, E. Sieben, J. Simaika, J. Šinžar-Sekulić, P. Skelton, K.G. Smith, The editors thank: E. Abban, H.M.H. Adliw, L. Agenbag, O. S. Smithies, D. Snijman, J. Snoeks, G. Soliman, A. Soumia, Akinsola, E. Akinyi, A. Alegro, M. Alexis, M. Ali, D.J. Allen, P. M. Sow, A.-S. Stensgaard, M. Stiassny, F. Suhling, K. Swart, Alves, A. Amadi, J.S. Amakye, S. Amara, G. Ameka, W. Amer, E. Swartz, S.I. Sylla, K. Tabu, M.I. Tattou, A.N. M. Tawe, S. K. Amoako, M. Angliss, C. Appleton, C. Archer, F.Y.K. Attipoe, Tchibozo, P. Tchouto, H.J. Temple, S. Terry, N. Thomas, S. A. Awaiss, A. Azeroual, D. Azevedo, S. Bagella, M. Baninzi, B. Todd, C.D. Touokong, C.D. Touokong, D. Tweddle, T. Twongo, Bayer, J. Bayona, G.M. De Bélair, S. Ben Saad, S. Benhouhou, D. Van Damme, E. Vela, J. Victor, J.-C. Vié, J. Viera, E. Vreven, L. Bennet, M. Benslama, M. Biague, C. Bigirimana, R. Bills, M.S.A. Warith, K. West, F. Wicker, F. Witte, A. Yahyaoui, R. x E. Bizuru, T. Bokhutlo, J. Boudot, L. Boulos, T. Bousso, Zaiss, and A.I. Zamba. INTRODUCTION Foreword

For millions of people throughout Africa, wetlands are we hope that decision-makers in Africa will now make | critical for livelihoods, providing vital supplies of water, food the right choices to develop their water resources in and materials as well as ecological services. Wetlands are, a sustainable manner whilst protecting and valuing FOREWORD however, suffering from high levels of degradation, and this biodiversity. is likely to increase dramatically over the next few decades as Africa strives to provide food, water and electricity to an The Ramsar Convention on Wetlands covers all aspects increasing population. of wetland conservation and wise use, recognising wetlands as ecosystems that are extremely important for In the most comprehensive assessment of its kind, 4,989 biodiversity conservation and for the well-being of human African freshwater species were assessed by close to 200 communities. A key commitment of Contracting Parties, or scientists over a six-year period for the IUCN Red List Member States, is to designate suitable wetlands for the of Threatened Species, including all described species List of Wetlands of International Importance (‘Ramsar List‘) of freshwater fi sh, molluscs, crabs, dragonfl ies and and ensure their effective management. This volume, and damselfl ies, and selected families of aquatic plants. The the data collated through the project, will greatly assist fi ndings from this new study give us a unique opportunity those parties to identify new Ramsar sites, to broaden to try to infl uence developers and governments when they their taxonomic coverage, and to help ensure the future are planning freshwater infrastructure projects, which are wise use of wetlands in general. I therefore commend still in their early stages in most of Africa. Until now we this report and the accompanying information to you, have not had the information we need about species and and I urge all those concerned with the management and the threats they face, but, armed with these IUCN Red List conservation of wetland resources, to read it and to use assessments, the extensive information upon which they this most extensive new data set in their future work to are based, and the overall fi ndings and recommendations, secure the future of Africa’s wetlands.

Anada Tiéga Secretary General The Convention on Wetlands Ramsar Secretariat Switzerland

xi Executive Summary

In this volume you will ﬁ nd the most up-to-date information globally threatened. This level of threat is relatively high on the distributions and conservation status of species in in comparison to other taxonomic groups in Africa (12% all inland water ecosystems across mainland continental of birds, 19% of mammals, and 26% of amphibians are Africa and the reasons behind their declining status. This threatened) and is predicted to increase dramatically unless

EXECUTIVE SUMMARY represents the most comprehensive assessment yet of the ecological requirements of freshwater species are given

| freshwater biodiversity at the species level for an entire much greater consideration in future development planning, continent. For managers, this information will assist in in particular for development of water resources. Major designing and delivering targeted action to mitigate and threats are identifi ed as loss and degradation of habitat minimise these threats. From a policy perspective, this associated with deforestation, agriculture and infrastructure information is fundamental to meeting national obligations development, unsustainable levels of water extraction, water under the Convention on Biological Diversity (CBD), the pollution from domestic industrial and agricultural sources, Ramsar Convention, and the Millennium Development the introduction of alien invasive species, sedimentation, INTRODUCTION Goals (MDGs), and will input to national-level conservation mining and subsistence use and trade. The majority of priority setting. Information on species status is particularly threatened species are found along the Mediterranean and important for Target 12 of the CBD: ‘…by 2020 the extinction Atlantic coasts of Morocco, Algeria and Tunisia, in Upper of known threatened species has been prevented and their and Lower Guinea, southern and eastern South Africa, and conservation status, particularly of those most in decline, in the Great Lakes in eastern Africa. This distribution largely has been improved and sustained.’ refl ects a combination of high levels of current development activity, and high pressure on water resources, relative to Biodiversity within Africa’s inland waters is both highly other parts of the continent. diverse and of great importance to livelihoods and economies. Africa is, however, about to embark upon an Major centres of species richness include the Great Lakes of unprecedented scale of development within its water and eastern Africa, rivers in coastal sub-catchments in western agriculture sectors. Such development is proposed by Africa, and sub-catchments that trace the course of major many as essential if Africa is to support and improve the rivers through the centre of the continent, primarily in the livelihoods of a population that is projected to double by Democratic Republic of Congo (D. R. Congo). A network 2050 to almost two billion people. Development activities of river and lake basins is identifi ed as potential Key are, however, not always sustainable or compatible with Biodiversity Areas (KBAs) most important for the protection species conservation in inland waters, the needs of which of threatened and restricted range species. Forty eight sites are poorly represented within the development planning are proposed as new Alliance for Zero Extinction (AZE) sites process. holding Critically Endangered or Endangered freshwater species in most urgent need of conservation action. These One commonly cited reason for the inadequate representation proposed AZE sites should form the focus of the most of biodiversity in development and environmental planning immediate conservation actions if species extinctions are is a lack of readily available information on freshwater to be prevented. species. To close this information gap, the IUCN Species Programme, in collaboration with its partners, conducted Africa’s freshwater species are recognised to be as important an assessment of the status (according to the IUCN Red today as they ever have been for supporting livelihoods. List of Threatened Species) and distribution of all described The most heavily used species are the fi shes and plants, species of freshwater fi shes, molluscs, odonates, crabs, with 45% and 58% of species harvested, respectively. and selected families of aquatic plants from across mainland Justifi cation for the conservation and sustainable use of continental Africa. A total of 4,989 species were assessed. these species should, therefore, be clear to all. With species information compiled for each of 7,079 river or lake sub-catchments, this represents a major advance Inland waters throughout Africa are poorly represented in knowledge for informing development actions at a scale within the existing protected areas network, which is largely appropriate for conservation management. The full data designated for the protection of terrestrial ecosystems set, including all species distribution fi les, is available on and species. Future conservation efforts must take greater the DVD accompanying this report and through the IUCN account of the upstream and downstream connectivity in Red List website (www.iucnredlist.org). freshwater ecosystems. For example, it is recommended that conservation efforts focus on the protection of upper Twenty one percent of all species assessed are threatened catchment areas, provision of adequate environmental within Africa. Ninety one percent of these species (4,539) fl ows, and the inclusion of rivers within protected areas xii are endemic to the continent and are, therefore, also rather than as the boundary markers. Integrated river INTRODUCTION basin management and systematic conservation planning the continent to be managed in a way that ensures their approaches are recommended, along with the establishment long-term sustainability and maintains their ability to adapt of additional river and lake basin authorities. to changing conditions that they may face in the future. With this in mind, a number of case studies have been Patterns of past and present species diversity suggest that conducted, as a key component of this project, to develop Africa’s freshwater species have been highly resilient to a series of ‘Good Practice Guidelines’ for the integration | climate and geological change over time. The challenges of biodiversity information within the environmental and facing African freshwater species are daunting, but the development planning processes. The recommendations EXECUTIVE SUMMARY continent’s climate history suggests that signifi cant resilience from these studies can be found at: www.iucn.org/species/ exists. With informed development and management, freshwater/ African freshwaters may see a new century with different but not unhealthy qualities. The IUCN Red List is one of the most authoritative global standards supporting policy and action to conserve species. Following the six years of work required to complete this We hope this analysis, based in large part on an assessment study, we recognise that the greatest challenge is to ensure of species Red List status, will provide new information and the knowledge gained is transferred to the relevant decision insights, which will motivate actions to help safeguard an makers and stakeholders, and that it is updated on a essential and valued resource for millions of people – the regular basis. This will enable freshwater systems across diversity of life within Africa’s inland waters.

Key messages

➊ The inland waters of Africa support a high diversity legislation, without catchment-based planning that of aquatic species with high levels of endemism. extends the designated control areas to the edge of Many of these species provide direct (e.g., fi sheries) the river catchment, impacts such as water pollution and indirect (e.g., water purifi cation and fl ood control) and invasive alien species will inevitably lead to benefi ts to people. The conservation and sustainable species decline. management of these species is essential to the ➎ Management of water resources must take account livelihoods and economies of Africa’s people. of the requirements of freshwater biodiversity. If ➋ Current levels of threat to freshwater species we are to conserve and continue to benefi t from the across Africa are high relative to other ecosystems, services provided by freshwater species we need with 21% of species threatened. Predicted future to manage water as a resource for both people and levels of threat, in particular due to development of freshwater biodiversity. This approach is encapsulated water resources, are expected to be even higher. This within the Environmental Flows concept, which aims is largely a result of: i) the high degree of connectivity to ensure that there is enough water to maintain within freshwater systems, such that threats like environmental, economic and social benefi ts. pollution and invasive alien species spread more ➏ rapidly and easily than in terrestrial ecosystems; and Lack of available information should no longer be ii) the rapidly increasing use and development of water given as a reason for inadequate consideration resources, with little regard to the requirements of the for development impacts to freshwater species. freshwater dependent species sharing the resource. The data made available here must be integrated within the decision-making processes in planning for ➌ Protected areas must be better designed to protect the conservation and development of inland water freshwater species. The knowledge collected during resources, ensuring all development projects have a this assessment can be integrated into conservation ‘Net Positive Impact‘. planning processes to ensure representation of the freshwater species. Currently, protected areas ➐ Species information remains very limited for many are rarely designed to protect freshwater species, parts of Africa. The Congo, Angola, the Ethiopian meaning that taxonomic groups that represent a Highlands, and Upper and Lower Guinea in particular, signifi cant portion of the total species and genetic have been identifi ed as priorities for future fi eld survey. diversity on the planet are being overlooked. Information on the status and distribution of aquatic plants needs to be greatly improved. ➍ Protected areas and other conservation action for freshwater species must be designed to employ ➑ Environmental Impact Assessments should the principles of catchment management. Even expressly require reference to the species data where species are identifi ed by species-driven sets made available through the IUCN Red List. xiii Chapter 1. Background

Dudgeon, D.¹ (Section 1.1); Paugy, D. and Lévêque, C.² (Section 1.2); Rebelo, L.-M. and McCartney, M.P.³ (Section 1.3)

1 Chair of Ecology and Biodiversity, School of Biological Sciences, The University of Hong Kong, Pok Fu Lam Rd., Hong Kong SAR, China. 2 Institut de Recherche pour le Développement (IRD), Le Sextant, 44, bd de Dunkerque, CS 90009, 13572 Marseille, Paris, France 3 International Water Management Institute, C/o ILRI-Ethiopia Campus, Wereda 17, Kebele 21, Addis Ababa, Ethiopia.

MAIN: Lake Ichkeul in Ichkeul National Park, Tunisia. A World Heritage Site impacted by dams, but taken off the ‘sites in danger list’ in 2006 due the progress of restoration work. © HICHEM AZAFZAF RIGHT: Fisherman searching for his nets in the Sanaga River, Cameroon. © KEVIN SMITH CONTENTS 1.1 Global status of freshwater biodiversity 4 1.1.1 Species diversity 6 1.1.2 Major threats to freshwater species 8 1.2 Situation analysis for Africa 13 1.2.1 General overview 13 1.2.2 Threats to freshwater ecosystems 16 1.2.2.1 Habitat transformation 18 1.2.2.2 Pollution 22 1.2.2.3 Impact of fi sheries 23 1.2.2.4 Invasive alien species 24 1.2.2.5 Climate change 25 1.2.3 The African Green Revolution and wetlands 25 1.3 Functions and values of freshwater ecosystems in Africa 26 1.3.1 Ecosystem functions 26 1.3.2 Ecosystem values 28 4 CHAPTER 1 | BACKGROUND human water right,itis far from clearhowthe waterneeds aside from abouttheimplications ofa legalconcerns demand islikely toexceedthesupplyfrom runoff. Thus, et al planetary limitsfor‘bluewater’runoff resources (Rockström (Alcamo are ofwaterstress causingdramatic changesinpatterns rapid shiftsinanthropogenic wateruseandwithdrawal than halfofglobalsurfacerunoff (Jackson relating toquantity. Humansalready appropriate more aboutwaterqualityareConcerns paralleledbythose around 5,000 child deathsattributabletocontaminatedwaternumber UNICEF 2008).Theresult isaparloussituation,where than 2.5billion)ofpeoplelackadequatesanitation(WHO/ ready accesstodrinkingwater, andperhaps40%(more population (approximately 0.9billionpeople)doesnothave water andsanitation.Asignifi imperative toensure thatallpeopleshouldenjoyadequate may beproblematic, itisimpossibletodisagree withthe recognition thataccesstocleanwaterisahumanright no votesagainsttheresolution. Whileimplementingthe human right.Somecountriesabstained,butthere were access tocleanwaterandsanitationisafundamental Nations (UN)GeneralAssemblyresolution declaringthat On 1August2010,122countriessupportedaUnited 1.1 fundamental right. Girls collectingwaternearBukavu,D.R.Congo.AccordingtotheUnitedNations,accesscleanandsanitationisa . 2009a,b)sothat,withincurrent humanlifetimes, biodiversity Global statusoffreshwater et al . 2008).We maybecloseto overstepping daily ©IUCN/INTUBOEDHIHARTONO (approximately 1.5millionannually). cant proportion oftheEarth’s et al . 2001),and Dudgeon 2010).Forinstance, ithasbeen estimatedthat Revenga species threatened withextinction(e.g., Loh freshwater ecosystems tendtohavethehighestportionof greater thantheirterrestrial ormarinecounterparts,sothat population declinesandlosses from inlandwatersare al for marineandterrestrial biodiversityloss(Rockström an extentthatwehave‘oversteppedplanetaryboundaries’ 2010). Theseexceedbackground extinctionratestosuch Ehrlich andPringle2008;Butchart undergoing anepidemicofhuman-causedextinctions(e.g., that itisnowmore evidentthaneverthatthebiosphere is of Biodiversity. Thisserved,inpart,tohighlightthefact because 2010wasproclaimedYear astheUNInternational right, themidpointof‘Water forLife’decadeisnotable Apart from thepronouncement thatwater isahuman DecadeforAction(2005-2015). Life’ International as amajorachievementoftheUN-designated‘Water for of theMillenniumDevelopmentGoalsintendedtostand people withoutaccesstocleanwaterandsanitationisone matter isurgent, notleastbecausehalvingthenumberof it fl remaining inriversthatsustainswetlandsandestuariesas required byecosystems;forexample,the‘wasted’water needs willhavemajorimplicationsforthesupplyofwater terms. Itneverthelessseemsobviousthatmeetingthese of burgeoning humanpopulationscanbemetinpractical . 2009a,b).Moreover, aconsensushasemerged that ows tothesea.Furthermore –andself-evidentlythe et al . 2005;Dudgeon et al et al . 2006;Strayer and . 2010;Mace et al . 2005; et al et . CHAPTER 1 | BACKGROUND 5 cient to slow the rates cient to slow the . 2002). To achieve this, reliable achieve this, reliable . 2002). To et al © KLAAS-DOUWE DJIKSTRA of population decline and species loss from fresh waters fresh loss from decline and species of population planning, management, One thing is certain: (Strayer 2006). the need to must address processes and decision-making and human environmental between trade-off balance the or risk further biodiversity declines, needs for water, functioning and consequential impairment of ecosystem is a need to In short, there impacts on human livelihoods. habitats as and other freshwater ‘legitimize’ rivers, lakes users of water (Naiman and distribution of freshwater information on the status needed as well as the ecological conditions biodiversity, an essential component of to sustain it, must become Provision planning and water management. environmental a major objective of this report. of such information is phrase ‘the Stiassny (1999) co-opted Marshall McLuhan’s to encapsulate the notion medium is the message’ in order due biodiversity faces unparalleled threats that freshwater subject to unprecedented to dependence on a resource these threats demands. Here, human and ever-increasing so explained, with further discussion on why they are are and what makes the biodiversity associated with severe, waters particularly vulnerable to human impacts. fresh two explanations for are there Beginning with the latter, waters. vulnerability of biodiversity in fresh the relative deal of biodiversity in the small is a great there Firstly, of the globe occupied by rivers, lakes and wetlands. area the in section 1.1.1. Secondly, This point is addressed make their environments of freshwater features inherent by wrought inhabitants especially susceptible to changes considered are humans. The consequences of this matter to the pandemic array of threats in section 1.1.2, where biodiversity is described. freshwater biodiversity that sustains them (Naiman and Dudgeon Dudgeon and them (Naiman sustains that biodiversity that current been expressed have even Doubts 2010). suffi strategies will be conservation . 2008) . 2008) et al ts to be derived . 2005, Strayer 2006; . 2005, Strayer . 2006). One estimate . 2006). One estimate et al in Europe – almost 40% sh et al Moreover, the value of fresh the value of fresh Moreover, . . 2000), and the recently-declared . 2000), and the recently-declared . 2010) et al et al sh and unionid mussels) are threatened threatened are unionid mussels) sh and ood-recession agriculture, and dry-season and dry-season agriculture, ood-recession . 1997) puts the value of ecosystem services . 1997) puts the value . 2000; Revenga . 2000; Revenga . 2007; Strayer and Dudgeon 2010). Freshwater Freshwater and Dudgeon 2010). . 2007; Strayer et al et al shing in the rapids of the Boyoma Falls, on the Lualaba River in D. R. Congo. et al sheries, fl sheries, . 2007) is emblematic of their imperilment. Evidently, of their imperilment. Evidently, . 2007) is emblematic (Costanza waters at USD6.6 trillion annually, by fresh provided and in 20% of the value of all ecosystem combined, ecosystems excess of the worth of all other non-marine smaller extent combined (USD5.7 trillion), despite the far subject to are of inland waters. While valuation estimates waters have fresh the general message that controversy, self-evident. At immense economic importance seems for directly least two billion people depend upon rivers of ecosystem services that can be characterised provision most simply as ‘food’, such as the benefi fi from grazing (Richter Taylor Taylor mammals among the most endangered cetaceans are on Earth (Reeves River dolphin (Turvey of the Yangtze functional extinction et al hotspots of endangerment. waters are fresh component species waters and their Degradation of fresh is a matter for grave concern, and services given the goods them (Dudgeon derived from as ecosystems in the future, waters is bound to increase and their goods and services stressed become more scarcer. human needs It will be a colossal challenge to reconcile of goods and provision for water without compromising functioning ecosystems and the from services that result 20% of the world’s freshwater fi freshwater of the world’s 20% 2007; Jelks Freyhof and USA (Kottelat and the reptiles, and freshwater as many amphibians – as well species macroinvertebrate freshwater and 10,000-20,000 crayfi (especially (Gibbons Wagenia fi Wagenia 6 CHAPTER 1 | BACKGROUND species, makingthem‘ultra-hotspots’ for globalbiodiversity. habitatscoveronly 0.8%oftheEarth’sOkavango Delta channel. Freshwater surface butcontainalmost10%ofalldescribed only contain onlyaround 0.01%oftheEarth’s waterandcover are confi total, thenasmuchone-third ofallvertebratespecies river dolphins,platypus)are addedtothisfreshwater-fi aquatic reptiles (crocodiles, turtles)andmammals(otters, quarter ofglobalvertebratediversity. Whenamphibians, – approximately 40%ofglobalfi more than13,000speciesare fi et al on Earth,around 126,000liveinfresh water(Balian the approximately 1.32millionspeciesthusfardescribed expected from thearea occupiedbyinlandwaters.Of demonstrates thatitisverymuchlarger thanwouldbe incomplete –globalassessment(Balian especially intropical latitudes,butarecent –albeit The biotaoffresh watershasyettobefullyinventoried, 1.1.1 Speciesdiversity global biodiversity(StrayerandDudgeon2010). remaining unglaciatedregions mustbe‘ultra-hotspots’for low biodiversity, itisevidentthatfresh watersinthe situated inrecently-glaciated regions thathaverelatively (see above).Giventhelarge proportion offresh waters explaining whytheyare alsohotspotsofendangerment for globalbiodiversity, andthisgoessomewaytowards al . 2006).Fresh watersasawholeare therefore a‘hotspot’

. 2008)–almost10%oftheglobaltotal.Ofthese, about 0.8%ofthesurface(Gleick1996;Dudgeon ned tofresh water. Yet surfacefreshwater habitats s (Lévêque sh sh diversity, andone et al et al . 2008) . 2008) sh et © JENSKIPPING fi known: between1976and2000,more than300new well-studied freshwater taxaasfi either threatened ortoopoorlyknowntoassess.Evensuch known tobethreatened, atotalof56%amphibiansare as DD.Whenthisfi invertebrates: 25%ofamphibiansare classifi This insuffi rises from 16%to38%. with theadditionofDDcategory, andforcrayfi threatened freshwater crabsincreases from 16%to65% considered tobeatrisk(IUCN2010);theproportion of or thatare toopoorlyknowntoassessadequately) are those forwhichtaxonomicstatusremains uncertain, if speciesclassifi dragonfl et al (Clausnitzer groups globally:dragonfl assessments havebeenmadeforonlythree invertebrate status oftheseanimalsisproblematic. IUCNRed List et al especially intropical latitudes(Dudgeon are particularlynoticeableamonginvertebrates, and improving, large gapsremain (Balian Although knowledgeoffreshwater biodiversity is sh species,approximately 1%ofknownfi . 2009),andcrayfi . 2008).Accordingly, determinationofconservation ies are threatened, theproportion risesto40% ciency ofknowledgeisnotconfi et al ed as‘DataDefi . 2009),freshwater crabs(Cumberlidge gure iscombinedwith thepercentage sh (IUCN2010).Whileonly14%of ies (asampledassessment) cient’ (DD:essentially, shes are incompletely et al et al . 2008).They . 2006;Balian ed byIUCN se, were shes, nd to ned s it sh CHAPTER 1 | BACKGROUND 7

ects . 2004), ect the ies and et al . 2005; Chapter . 2006), and this diversity) between et al B et al . 2002; Benda ect the degree of isolation ect the degree et al . 2008). The high endemism results . 2008). The high endemism results uence patterns, gradients, or stream et al . 2005, Dudgeon . 2002), because there can be large ‘stream ‘stream can be large . 2002), because there et al et al ies (Balian contributes to regional species richness. contributes to regional Local endemism and high species turnover refl species to disperse through limited ability of most freshwater river to river along or to migrate from landscapes terrestrial arrangement of riverine the hierarchical the coast. Moreover, habitats means that the populations and communities connected to – or isolated differentially they harbour are (Fagan – each other from networks that depend on with abilities to disperse through the vagaries of confl barriers such as waterfalls. Geographic of the presence refl distance may appropriately which distance, stream whereas habitats, among terrestrial than straight-line distance, refl is often much larger habitats. Thus, among stream of isolation the degree tend to be isolated habitats for fully headwater streams geographically proximate aquatic species, even if they are (Gomi several major groups, including decapod crustaceans, several major groups, caddisfl molluscs and aquatic insects such as mayfl in considerable species turnover (= is cichlid basins or catchments. An outstanding example of hundreds lakes, where diversity in African Rift Valley found (Thieme endemic species are The fundamental point is that because 3 in this report). water bodies tend not to be of high species turnover, to their faunal assemblages ‘substitutable’ with respect (Revenga cation . 2008). . 2006), et al et al . 2000). More strikingly, strikingly, . 2000). More et al . 2000; Brooks . 2000; Brooks et al rmation that terrestrial and freshwater and freshwater rmation that terrestrial sh species (Sverdrup-Jensen 2002), more 2002), more sh species (Sverdrup-Jensen © MCKAY SAVAGE © MCKAY . 2008). Regional discovery rates of new freshwater freshwater . 2008). Regional discovery rates of new rst attempt at mapping global freshwater ecoregions and ecoregions rst attempt at mapping global freshwater despite the evidence that they are highly threatened and highly threatened despite the evidence that they are the lack of any confi hotspots overlap (Strayer and Dudgeon 2010). waters so rich in biodiversity? A few fresh Why are geographic ranges, but have large species freshwater habitats has led to the of freshwater the insular nature evolution of many species with small geographic ranges, often encompassing just a single lake or drainage basin (Strayer 2006; Strayer and Dudgeon 2010). High levels of local endemism and species richness seem typical of This is an important development, as previous global This is an important development, as previous freshwater ignored biodiversity assessments have largely species (e.g., Myers formally described or resurrected from synonymy each from formally described or resurrected year (Stiassny 1999; Lundberg 40% of the global total of 6,696 amphibian approximately two decades species has been described during the last likely are 2010). The equivalent proportions (AmphibiaWeb invertebrates, with substantial for freshwater to be more (Balian tens of thousands of species awaiting description et al Investigations in the Mekong species also vary widely. may support as drainage, for example, suggest that it many as 1,700 fi total of earlier estimates. Identifi times the than three richness particularly high freshwater that support of areas the and realm, for the terrestrial has lagged behind efforts fi (Abell recently hotspots was unveiled relatively A polluted canal in Chennai, India. Pollution of waterways from urban centresA polluted canal in Chennai, India. Pollution is a major threat to freshwater systems and their biodiversity. 8 CHAPTER 1 | BACKGROUND consequences willbe furthercompounded byglobal interactions between thesethreats or stressors, andtheir example (Hecky impacts ofNilePerch inLakeVictoria beinganegregious present anadditional threat tofreshwater biodiversity–the non-native oralienspecies(reviewed byStrayer2010) al global problem (reviewed byAllan addition, overexploitation offi for organisms (reviewed byDudgeon humans inwaysthatcompromise itsvalueasahabitat may beextracted,diverted,containedorcontaminatedby arises from thefactthatfresh wateris a resource that notion thatthevulnerabilityoffreshwater biodiversity ‘The mediumisthemessage’effectively conveysthe 1.1.2 Majorthreats tofreshwater species by humanactivities,asdescribedbelow. contribute toitsvulnerabilitythemanythreats generated Thus, thefeatures generatingfreshwater biodiversityalso local extinctionevents(Cook of recolonization orrecovery followingperturbationsor (through evolutionofendemism),butalsoinfl and/or isolationoffresh waterscancontributetorichness overland dispersalremains. Thehierarchical architecture lakes orotherstanding-waterbodies,buttheproblem of appropriate measure ofthedegree ofisolationamong distances’ betweenthem.Geographicdistanceisamore biodiversity andimpactingthewatersuppliestoKunmingCity. Algal bloomsinLakeDianchi,China.Industrialanddomesticpollutionhasledtotheeutrophicationoflake,threatening . 2006;Wong et al et al . 2007),whileintroductions ofexotic . 2010).There willbecomplicated et al shes andotheranimalsisa . 2007;Lake et al . 2005;Dudgeon et al uences rates uences et al . 2006).In . 2007). et ©GREENPEACECHINA as ‘receivers’ (Ward 1989).Forexample, pollution from so thatfl affected byupstream processes, includingperturbation, downstream assemblages instreams andriversare complex direct and indirect pathways.Furthermore, and catchmentconditionimpacts biodiversityviamultiple and riversare landscape ‘receivers’ (Dudgeon that are associatedwith biodiversitylosses.Inshort,lakes in eutrophication, toxicalgalblooms,fi extinction overlongertimescales.Pollutantsmayresult lethal effects andphysiologicalimpairmentthatmaycause biota bypoisoningandhabitatdegradation,sub- runoff andsoon,leadingtodirect patterns mortalityof can leadtosedimentation,pollutionbynutrients,changed areas. Landtransformationforagriculture orurbanization very signifi diversity oflakes,streams andriverstobedetermineda biodiversity isaresult ofthetendencyforintegrityand The multiplicityofhumanimpactsonfreshwater (Olden species concerned species are particulartolocalcircumstances andthe are at-riskduetooverfi be atgreatest riskofextinctionglobally, buttheformer largest andsmallestspeciesoffreshwater fi and consequentialextinctionrisks.Forexample,boththe predict likelyoutcomes(Moss2010;Ormerod runoff andprecipitationThismakesitdiffi patterns. climate changeleadingtorisingtemperatures andshiftsin owing-water habitats are ‘transmitters’ aswell cant extentbytheconditionoftheircatchment shing, whilethreats tothesmallest et al . 2007). sh killsandsoon sh appearto et al et al cult to cult . 2010) . 2006) CHAPTER 1 | BACKGROUND 9

ow ow ows. hidrovias oods and ow regime . 2007). Some ow conditions . 2005). Another et al et al oods and droughts) oods and droughts) . 2002; Matthews and Marsh- ow variability as undesirable or – sh and other organisms, whereas whereas and other organisms, sh et al . 2007). The consequences of the of the . 2007). The consequences et al ve times the volume of all the world’s rivers ve times the volume of all the world’s uxes are also typical of biologically-diverse also typical of biologically-diverse uxes are . 2007). Even species that occur in multiple lake lake occur in multiple that Even species . 2007). et al ed by the construction of dams (e.g., Nilsson ed by the construction of . 2005), and the interaction of this anthropogenic . 2005), and the interaction of this anthropogenic uvial navigation as industrial waterways or waterways industrial as uvial navigation ciency throughout drainage networks (especially drainage networks ciency throughout ow is problematic is that all fresh waters are spatially waters are is that all fresh ow is problematic for fl (Wong (Wong locally- by endemic, may be represented or river systems or individual water bodies lineages within adapted genetic units that unique taxonomic These represent drainages. genetic and mixing of due to interbreeding will be lost interbasin water transfers can accompany lineages that (Moritz 2002). with impacts can be transferred It is a paradox that effi – the complex yet – as mentioned above downstream), such networks and the isolation of water of architecture from and recovery resilience bodies tends to constrain impacts (Lake greatly waters are of fresh nature fragmented and insular magnifi et al network fragmentation and the geometry of the drainage aquatic the persistence of threatened affect can strongly species (e.g., Fagan impacts Matthews 2007). The extent of such dam-related of dam-based is very substantial: a global overview half (172 out of that over revealed rivers impacts on large fl by fragmentation, with the greatest affected 292) were (Nilsson River in the Volta regulation of global fl indication of the extent of human alteration the over 10,000km3 of water, is that dams retain regimes equivalent of fi trap 25% 2000), while reservoirs (Nilsson and Berggren the oceans it reaches of the total sediment load before (Vörösmarty and Sahagian 2000). of why alteration and regulation A fundamental reason fl variability in and temporally dynamic systems, exhibiting or other aspects of water regime, inundation discharge, on diurnal, as well as light and temperature, seasonal and variability in sediment timescales. Temporal inter-annual and nutrient fl and interact with fl ecosystems, freshwater (including disturbances such as fl to maintain habitat diversity and ecosystem processes, persistence and richness of native enhance and thereby species (Bunn and Arthington 2002; Poff adapted to ephemeral or intermittent of these species are for part of the year only, is present water systems, where inundation or fl perennial require others whereas and/or associated Seasonal peaks in the hydrograph or turbidity may represent changes in temperature cues for fi reproductive only during low-fl some species may recruit 2004). and Poff (Bunn and Arthington 2002; Lytle fl Humans tend to treat (for example, fl – disastrous cases in extreme modify or engineer freshwater and therefore drought), ow shes . 2010). Further ooding (Lytle and ooding (Lytle et al shing have caused . 2002). Migratory fi et al ow of water. Examples include ow of water. oodplains by limiting or severing their oodplains via seasonal inundation during high- oodplains via seasonal shes or shrimps that breed in estuaries, thereby resulting resulting in estuaries, thereby or shrimps that breed shes fry with limited genetic variability sh or hatchery-reared ow periods (Ward 1989). Many riverine or wetland species 1989). Many riverine ow periods (Ward upstream is transmitted downstream, thereby spreading spreading thereby downstream, transmitted is upstream Disturbances intact reaches. to otherwise impacts potential can also be transmitted riverine biodiversity that threaten against the fl upstream migration of upstream dams that impede downstream fi in headwaters of whole assemblages in extirpation and overfi (Pringle 2001). Dams losses of salmon runs in rivers along declines or complete America, leading to reductions the west coast of North marine-derived nutrients with major in ‘uphill’ transfer of in production and riparian consequences for in-stream headwaters (e.g., Gende degrades fl regulation in turn,connection with the river channel which, impacts and of aquatic species (Lytle migration and reproduction 2004). Changes to riparian zones or river banks Poff transfers of energy disrupt food webs and the reciprocal and aquatic habitats and nutrients between terrestrial (Nakano and Murakami 2001; Fausch impact can be attributed to susceptibility to anthropogenic waters ground the fact that exchanges between surface and fundamental for maintaining the integrity of freshwater are surface and ground ecosystems although, typically, 1989). (Ward managed as separate resources waters are freshwater connectivity between the inherent In summary, that catchments ensures bodies and their surrounding can originate well beyond lake or to biodiversity threats connectivity allows river banks, and within-river hydrologic and upstream impacts to be transmitted both downstream the relatively from (Pringle 2001). This is markedly different of most human impacts in terrestrial localized effects landscapes. One category of human impacts that is peculiar to inland to species endemism at the basin waters is the threat scale posed by transfer of alien species among river systems (e.g., deliberate stocking with non-indigenous fi Strayer 2010), and the particular threat by – see review and mixing of formerly isolated ‘introduction’ arising from interbasin water large-scale through about species brought of rivers planned or ongoing transfers and interconnection in China, India, southern For Africa and the Neotropics. instance, the La Plata River System is listed as one of on account of plans the ten rivers most at risk globally, Orinoco and Amazon Rivers to connect the Paraguay, (alewives, eels, salmon and shad) have likewise declined (alewives, eels, salmon of abundance in historic levels from by as much as 98% and Waldman (Limburg seaboard rivers along the Atlantic 2009). the linkage between rivers Further complexity arises from and their fl fl adapted to and dependent on such fl are 2004). Levee construction, channelization and fl Poff 10 CHAPTER 1 | BACKGROUND and rivers isaprerequisite for protecting freshwater Maintaining the dynamicandvariablenature ofstreams habitat (Bunn and Arthington2002;Poff fl reproduction (Lytle andPoff 2004), a‘fl As wellasleadingtoaloss of hydrographic cuesfor bodies toincrease predictability andcontrol variability. water, theequivalentoffi Riano Dam,Spain.Anindicationoftheextenthumanalteration ofglobalfl ows limitsthefl uvial disturbanceneededtorejuvenate ve timesthevolume ofalltheworld’s rivers. attening’ ofpeak et al . 2007). e-fl regionally-specifi While newandinnovative strategiestodevelop paradigm aimedatcontrolling hydrological variability. challenge giventhecontextof aresource management biodiversity (Poff 2010; Poff ows) are beingdeveloped(Arthington © FELIPEGABALDON over10,000km³of isthatdamsretain ow regimes et al . 2010), including holistic methodsused in c environmental waterallocations(or et al . 1997),butpresents aformidable et al . 2006, CHAPTER 1 | BACKGROUND 11 . et et sh et al . 2009) (see also Chapter shing leading to reductions in leading to reductions shing et al . 2005b; Parmesan 2006; Heino . 2010). The term ‘ecosocial anomie’ et al uenced nutrient transfer, food webs and uenced nutrient transfer, . 2005). Hydrological changes, caused . 2005). Hydrological et al ows is evidence of growing awareness of awareness of growing evidence ows is shers can be affected by baseline shift within shers can be affected et al (IPCC 2007), and will have profound oods and droughts ect conditions in the intermediate and distant past: i.e.,ect conditions in the intermediate and distant . 2009). Warmer temperatures will have direct metabolic will have direct temperatures . 2009). Warmer . 2004). Allowing for some differences in scale or intensity in scale or intensity differences . 2004). Allowing for some al and could impact organisms on ectothermic effects thermal tolerances (e.g., Poff species with narrow 2001; Allan by changes in the amount and timing of precipitation, evapotranspiration, and glacial melting, may also be and severity in the frequency associated with an increase of fl Temperature biodiversity. implications for freshwater may be smaller than those further in the tropics increases (IPCC 2007), but this does not mean the equator from 8, section 8.6.1, this volume). Evidence for impacts on in the form of warmer has appeared waters already fresh and changes in periods of ice cover, shorter temperatures, animals the geographic ranges or phenology of freshwater by Allan (reviewed Research on e-fl on Research importantly, more and, biodiversity to freshwater threats it must be them. However, address an attempt to represents not are biodiversity that declines in freshwater emphasized have been waters on fresh phenomenon. Impacts a recent of places and, in the case centuries in some sustained over 4,000 persisted for over rivers in China, have many major fi freshwater Declines in European years (Dudgeon 2000). have been attributed to a combination 1000AD around from nutrient increased intensive agriculture, of siltation from dams, introduction of mill loads and pollution, proliferation overfi of exotic species, and 2005; Barrett 1996, (Hoffmann mean size and abundance al that threaten much the same factors of action, these are Historical losses of salmon, biodiversity today. freshwater American rivers are North from beaver and unionid mussels certain to have infl (Humphries and Winemiller 2009; dynamics organic-matter well 2009). Such impacts occurred and Waldman Limburg any formal stock assessments, giving rise to the before that conditions in the immediate past false impression refl the extent deception, and thus a tendency to underestimate (Humphries of human impacts, due to a ‘shifting baseline’ is not just aand Winemiller 2009). The shifting baseline species and charismatic large matter of historic interest: exploited by fi species are the span of a human generation; when these rapidly basis, they are on a fairly regular not encountered (Turvey forgotten in expectations has been used to describe this breakdown waters, and in fresh of what species should be present and (Limburg in their preservation societal loss of interest 2009). Waldman to present Human-caused climate change is certain and now biodiversity, to freshwater threat a profound dominates discussions about the fate of ecosystems marine, and and conservation planning in the terrestrial, (Heino realms freshwater cation of dam operations to mitigate their impacts cation of dam operations to have been reported (Richter and Thomas 2007; Olden (Richter and Thomas 2007; Olden have been reported and Naiman 2010). Asia and Africa that attempt to strike a balance between (King and Brown protection development and resource scales 2006, 2010), their implementation at appropriate will be challenging. Nonetheless, some successes with modifi 12 CHAPTER 1 | BACKGROUND forced more than20millionpeoplefrom theirhomes. forced more monsoonrainfallin2010that Flooding acrossvasttractsoflandinPakistan'sSindhprovince,sixmonthsonfromtheextreme where drainage basinsare oriented east–west. One compensatory movements northorsouthare notpossible limited inhuman-dominated environments. Moreover, and amphibiansmightfi terrestrial landscape (Dudgeon 2007).Evenfl fully aquaticspeciesthatcannot movethrough the altitudes are oftennot possible,especiallyforthemany into coolerhabitatsfurtherfrom theequatorortohigher to climatechangebywayofcompensatorymovements Given theinsularnature offreshwater habitats, adaptations and reduced adultfi temperatures willleadtosmallersizeatmetamorphosis invertebrates) hastheimportantimplicationthatwarmer growth andbodysizeinamphibians(andmanyaquatic The inverserelationship betweentemperature during close totheiruppertolerancelimits(Deutsch affected byrisingtemperatures, sincethey mayalready be fi that tropical ectotherms(‘cold-blooded’animalssuchas sh andamphibiansaswellinvertebrates)willbeless tness (e.g.,Semlitsch nd theirdispersalopportunities et al ying insects ying et al . 1988). . 2008). © DFID–UKDEPARTMENT FORINTERNATIONAL DEVELOPMENT thermal range(Dudgeon2007;Hoegh-Guldberg species from warmingwaterbodiestohabitatswithintheir would betranslocationoraidedmigrationofthreatened conservation initiativethatcouldhelpaddress thisproblem and water diversionsto enhancewater securityforpeople Palmer solutions tomitigate theseproblems (Dudgeon2007; life andproperty, effects thatwillencourageengineering exacerbate water-supply shortagesandthreaten human be asstrong orevengreater. Climatechangewillcreate or to thisclimatechangecouldgive risetoimpactsthatwill biodiversity overthecomingcentury, buthumanresponses Climate changewillhavestrong direct effects onfreshwater with alienspecies(StrayerandDudgeon2010). or leadingtotheecologicalimpactsoftypeassociated climate change),andposetheriskoftransferringdiseases for onlyatinyfractionoffreshwater speciesthreatened by detailed informationaboutthespecies(whichisavailable Such actionswouldbecontroversial andcostly, requiring et al . 2008),including newdams,dredging, levees, et al . 2008). CHAPTER 1 | BACKGROUND 13 et oodplains of ed – extinction ed – extinction . 2006), and demonstrate how human how human and demonstrate . 2006), et al . 2010). Extinctions are likely to continue over the next to continue over the likely are . 2010). Extinctions al resulting of actions taken now, regardless few decades, – and unquantifi unredeemed large a from by habitat instance, be incurred ‘debt’. This could, for populations that are the viability of fragmentation reducing (Matthews of dwindling to extinction now in the process 2007; Strayer and Dudgeon 2010). and Marsh-Matthews In June 2010, 85 so. but not entirely The outlook is bleak, Science to establish an Intergovernmental nations agreed and Ecosystem Services policy platform on Biodiversity is intended to The IPBES Korea. (IPBES) in Busan, South Change Panel on Climate the Intergovernmental mirror on declines in(IPCC), and will work to integrate data with thebiodiversity and degradation of ecosystems them (Larigauderie to reverse government action required global- and Mooney 2010). The IPBES will co-ordinate on the status and trends of research scale peer reviews and provide of biodiversity and ecosystem services to and policy recommendations reports ‘gold standard’ governments. of the IPBES will require Formal establishment but the initial by the UN General Assembly, agreement was an important initiative for the Internationalagreement a conduit will also provide The IPBES of Biodiversity. Year such as this can achieve wider currency by which reports inform conservation action. and thereby 1.2 Situation analysis for Africa 1.2.1 General overview the muddy waters of the Congo River spilling From to the vast fl into equatorial swamp forests the deep the Inner Niger Delta or Okavango Delta, from and shallow lakes of the Rift lakes in eastern Africa to of the Namib or Sahara deserts, from ephemeral streams basins to the mountain and the equatorial and perennial or South Africa, the inland coastal rivers of the Maghreb diverse. incredibly waters of Africa are has been For a long time, the African environment threats. serious anthropogenic from preserved relatively presently This is no longer the case, and several threats with and biological resources, both the hydrological affect biodiversity in some areas the rate of loss of freshwater to be high. For example, Begg (1988) suspected already showed that in the Mfolozi catchment of Natal, South had been lost Africa, 58% of the original wetland area by the mid-1980s. Another well-known example is from at the beginning of the 1980s, some where, Lake Victoria, to have reported 70% of the endemic cichlid species were and Lowe-McConnell 1994). (Worthington disappeared extent to which current practices are unsustainable unsustainable are practices current to which extent (Dudgeon have of inland waters and impairment exploitation management (Ormerod our best attempts at outpaced et oods. . 2008; Strayer et al . (2009) assert, it is evident that we must also have far oods associated with rainfall extremes) when there may when there oods associated with rainfall extremes) and agriculture and provide protection from fl from protection and provide and agriculture impetus to install new is increasing In addition, there dependence along rivers to reduce facilities hydropower needs. The global energy on fossil fuels and meet growing will ecological impacts of such engineering responses impacts of climate change, especially if magnify the direct to disasters (such as severe in response initiated they are fl reviews environmental be a temptation to circumvent need for project of the urgent because and regulations implementation (Dudgeon 2007; Palmer al would have been sustainable exceeded whatever margins Indeed, the extent of the biodiversity. for freshwater biodiversity that have declines and losses of freshwater indicator of the a reliable been documented is probably and Dudgeon 2010). planetary boundaries for transgressed If we have already biodiversity loss, as Rockström and marine terrestrial 14 CHAPTER 1 | BACKGROUND (redrawn from Beadle1981). Figure 1.1.ThegeneraltopographyofAfrica,showing theapproximate divisionbetweenHighandLowAfrica have a long evolutionary historycompared to temperate Africa isanancient continent,where freshwater systems Geology andgeography characteristics, biodiversityand values. the baselinedataonwetlandarea, distribution,seasonality, As aresult, updatedinventories are necessarytoimprove Africa,arethe CongoBasinandeastern stillpoorlyknown. been reasonably wellinvestigated,whileothers,suchas Africa,have Some areas,andwestern suchassouthern knowledge offreshwater systemsisquiteheterogeneous. aquatic faunaisstillratherpoorlyknown.Moreover, our with humanhealth(e.g.,diseasevectors),theAfrican (for example,fi Except forafewgroups ofeconomicallyimportantspecies shes andshrimps)orspeciesassociated most ancient lakesinthe world,suchas LakeTanganyika Valley, aresult oftectonicevents, includessomeofthe is mainlyover1,000m abovesealevel.Thefamous Rift sea level.Incontrast,HighAfrica tothesouthandeast sedimentary basinsandupland plainsbelow600mabove 1981) (Figure 1.1).Low Africaislargely composedof can broadly bedivided intoLowandHighAfrica(Beadle Taking intoaccountthecurrent relief, theAfrican continent events, combinedwiththeimpactsofclimatechanges. continent hasbeendeterminedlargely bythesegeologic present distributionoffreshwater species across the activity, mainlyduringthepast20millionyears.The that haveresulted from landtransformation, bytectonic zones. Theyoffer ahighnumberandvarietyofhabitats CHAPTER 1 | BACKGROUND 15 ned a biogeographic . 2008). estimate ora. A preliminary et al nal set of 11 bioregions and nal set of 11 bioregions conservation biology sh, freshwater ned for application to Africa’s freshwater freshwater ned for application to Africa’s nities at higher taxonomic levels. Based on nities at higher taxonomic levels. Based on © KLAAS-DOUWE DJIKSTRA . 2005). Furthermore, they defi . 2005). Furthermore, province, or bioregion, as a complex of ecoregions that as a complex of ecoregions or bioregion, province, history and so often have a similar biogeographic share affi strong existing knowledge of fi a fi and aquatic entomology, for Africa was delineated (Figure ecoregions 79 freshwater FADA provides a global overview of genus and species- a global provides FADA inland waters at level diversity of animals and plants in the information the continental level. Within continental Africa, biogeographical two large has been compiled across south of the Sahara and Madagascar belong to areas; to the north while the Maghreb region, the Afrotropical combining the Through region. belongs to the Palearctic it is possible to obtain an information for these two regions, freshwater estimate for the diversity of the aquatic African studied) and fl fauna (58 groups species occur in that at least 16,789 freshwater proposes and alone (including Madagascar region the Afrotropical islands) (Balian other offshore Main ecological zones in Africa attempts to delineate geographic have been several There Africa based on the origins and similarities of across areas of these is based on the aquatic fauna. The most recent (Olson and Dinerstein 1998), which concept the ecoregion has been refi of land or water containing a area ecosystems as a large distinct assemblage of natural communities and species, the original extent of whose boundaries approximate major land use change (Thieme natural communities before et al Fish catch in a bucket, caught in the Lower Itimbiri River in D. Fish catch in a bucket, caught in the Lower Asia and South R. Congo. Fish diversity in Africa rivals that of America. 2008). oods. Little is known et al. Jg\Z`\jèk_\Jgfkc`^_k

fZbf]]i\j_nXk\iZiXYjèCXb\ sh fauna (Lévêque and Paugy 2006), ows and lake levels. Evidently, these large these large ows and lake levels. Evidently, , this volume). sh and invertebrates (Worthington and Lowe- and (Worthington sh and invertebrates uence the ecology of the continent and its uence the ecology of the continent . 2008). In addition, the existence of diverse species 8le`hl\jg\Z`\jÕ sh diversity, reported to include more than 3,000 species, to include more reported diversity, sh a special additional ’living fossils‘ provides ocks and relict One to other continents. sh families in Africa as compared ocks of fi Æ KXe^Xep`bX1 X df[\c ]fi jkl[`\j\okèZk`fe è \mfclk`fe Xe[ Climate and hydrology importance in determining the The climate is of utmost Africa, systems. Within tropical distribution of aquatic (i) equatorial: hot major climatic types prevail: three hot with rainy seasons; (ii) tropical: and humid with two arid. In the north hot and subtropical: summer rain; and (iii) with climate types also occur, and south, Mediterranean rains. The amount of rain and arid summers and winter within greatly the year varies its distribution throughout in rainfall Africa. Both within and between year variations determine river fl spatial and temporal patterns in rainfall and temperature infl greatly aquatic organisms. 30 central Africa almost entirely covered Rainforest between 16 million years ago, but savannas expanded years remained and 8 million years ago. The last million than now for 80- with less rainforest changeable and dry, the continents 90% of the time. Just 12,000 years ago, While the area. only 10% of the current covered rainforest 8,000 years Sahara was dotted with swamps and lakes arid phase culminated only 2,500- ago, the most recent less rainfall Africa still receives present. 2,000 years before seasonal and and what does fall is more than other tropics, and fl droughts with frequent irregular, such constant about the long-term consequences of aridity and seasonal fauna. Increasing shifts for freshwater instance, could variability within and between years, for areas. affected extinction rates in more lead to increasing biodiversity Freshwater of geographic, climatic the combined effects Altogether, and topographic factors have given rise to a high diversity freshwater and assemblages. Africa’s species of freshwater fi than 3,600 species known) and rivals that of Asia (more than 4,200 species known) (Lévêque South America (more et al fl value to the African fi and there are an unusually high number of primary division are and there fi phylogenetic diversity may be for this exceptional reason that the African continent has not been totally submerged, at least for the last 600 million years (Lévêque 2001). Information on the diversity of other taxonomic groups Animal Diversity the Freshwater is compiled through (Balian project Assessment (FADA) (12 million years old (Fryer 1991)) and Lakes Malawi and and Lakes Malawi 1991)) and old (Fryer years (12 million species to unique given rise has evolution where Victoria, fl McConnell 1994) (also see McConnell 16 CHAPTER 1 | BACKGROUND Figure 1.2.Freshwater ecoregions ofcontinentalAfrica. rainfall to runoff, itisclearthat large areas ofAfricaface rate (around 65%ofrainfall),and lowconversionof Given theunpredictable rainfall,very highevaporation species; pollution;andoverexploitation. extraction andhydrological disruption;invasivealien scale, including:habitatloss or transformation;water ecosystems are recognised tooperateatthecontinental A numberofanthropogenic threats to freshwater 1.2.2 Threats tofreshwater ecosystems and Paugy2006;2010). fl corresponding totheGreat Lakesendemic fauna(species 1.2). Twelve provinces andseveral’hotspotprovinces‘ ocks) were delineatedforfi sh (Skelton1994;Lévêque and Richter2003; resulting inimpairmentoftheirecologicalfunction(Postel as dams,fl large wetlands havebeenaffected bydevelopments,such to meettheincreasing demandforwaterandpower, many demands forreliable andsafewatersupplies.Inaneffort water managementdiffi with an estimated 85% ofAfrica’s total water withdrawals total waterconsumption from rivers, lakesandaquifers, agriculture orindustryaccountsfor alarge partofthe (Darwall Africa particularly trueforareas andsouthern ofnorthern being lowered andwetland areas disappearing;thisis they canbereplenished, resulting inthewatertable to provide irrigation for agriculture, atafasterratethan 2006). Groundwater reserves are beingused,mostoften et al ood control measures, ordirect abstraction, . 2009;Garcia culties insatisfyingthegrowing et al et al . 2003;Dudgeon . 2010).Abstractionfor et al . CHAPTER 1 | BACKGROUND 17 .

et al shes in c impacts to other taxonomic groups are are c impacts to other taxonomic groups oodplains, with a consequent impact on oodplains, with a consequent cation is often inadequate or non-existent (Saad and their fl domestic sewage Pollution from associated biodiversity. in many a major problem and industrial facilities is already for water required the infrastructure parts of Africa where purifi 1994; Lévêque 1997; UNEP 2004). in more treated are each of these threats The impacts from a discussion of impacts on fi through detail below, Specifi particular. 3 to 7. in Chapters also covered © MANGUS FRANKLIN Polluted water from a leather tannery in Addis Ababa mixing with the Akaki river. Industrial water pollution is major threat to Polluted water from a leather tannery in Addis Ababa mixing with the Akaki river. freshwater biodiversity in many parts of Africa. directed toward agriculture (FAO 2010a), and about one- (FAO agriculture toward directed estimated to be under agricultural area of its surface third compound existing 2010b). To land use in 2000 (FAO Africa is poised to embark on water resources, pressures scale of development within its on an unprecedented in irrigated land a 100% increase targeting water sector, 7% to 25% of total hydropower from and an increase area, by 2025 (Economic Commission for potential captured these agricultural lands from runoff Africa 2003). Water brings sediments, nutrients and pesticides that can functioning of wetlands the effective or reduce destroy 18 CHAPTER 1 | BACKGROUND 2005; biodiversity are welldocumented (e.g., Nilsson The impactsof large damsonecosystemfunctions and production (Figure 1.4). including, irrigation,watersupply (domesticuse)orfi have beenestablishedtomeet otherwaterdemands More recently, probably thousandsofsmallerreservoirs the mid1950s,onlarge riversandforelectricitysupply. hydroelectricity. Mostofthelargest damswere built after industrial andminingwatersupply, crop irrigation and built forarangeofpurposes,includingdomestic, proposed. Large damsorsecondaryhavebeen many more are eitherunderconstruction orhavebeen Africa(FigureMaghreb, andwestern 1.3), southern Africa hasalarge numberofdams,particularlyinthe Dams (waterfragmentation) 1.2.2.1 Habitattransformation from FAO, Aquastat2010a). Figure 1.3.Locationof135largedams(

McAllister et al. 2001;McCartney et al r 500,000m³)and1,072secondarydams(

of ecosystem valuesdoes occur, through thecreation ecosystems. In somecases,however, enhancement cases, tosignifi more negativethanpositive andtheyhaveled,inmany Dams 2000):’Onbalance,the ecosystemimpactsare the World CommissiononDams (World Commissionon up through oneofthe mainconclusionsoftheReport of itsconstruction.Thenature oftheseimpactsissummed taken before buildingadam,andinthesubsequentdesign treated ascoststosocietyandbefactored intodecisions The ecosystemimpactsfrom damsshould,therefore, be many hundreds ofkilometres from thesiteofdam. quality. Suchchangesinecosystemfunctionscanbefelt changes tothefl for someaquaticspecies.Downstream impactsinclude submerged habitats,andblockthemigrationpathways and sedimentsalongrivers,permanentlydestroy upstream constitute obstaclestolongitudinalexchangeofnutrients cant andirreversible lossofspecies and ow regime, water temperature andwater CHAPTER 1 | BACKGROUND 19 . et al . 2005). et al ow. © INTERNATIONAL RIVERS © INTERNATIONAL An aerial view of the Inga Dam on the Congo River which, due to lack of maintenance, was running at an estimated 20% capacity in 2006. was realised that Grand Inga Dam would create a lake so that Grand Inga Dam would create was realised both Kinshasa and Congo that it would submerge large environmental Brazzaville and would cause ’irreversible at the river mouth‘. harm to all living matter and organisms disruptions to the Congo River The potentially catastrophic to of threats and the Atlantic Ocean interaction, in terms The also realised. global carbon and water cycles, were empty into Amazon, Orinoco and Congo rivers, which of the world the Atlantic, combined contribute 15-18% carbon inputs (Coynel total organic ocean’s Inga III and IV were proposed in 2009 as ’run of the river‘ proposed Inga III and IV were that do not disrupt the river fl structures For a few major impoundments in Africa, the sequence after damming has been studied (e.g., Jackson rst culty of coping shing and recreational shing and recreational nal 96m to sea level, the channel ow through a bank of electricity generators ow through cant falls or rapids close to its mouth. Furthermore, its mouth. Furthermore, cant falls or rapids close to ow (after the Amazon), but it is the only major river with as the river drops its fi as the river drops bend – this makes a 180 degrees and its course narrows is Inga Falls. The Grand Inga scheme would divert the this a channel cutting across lower Congo River through bend, to fl river to its natural course. Grand Inga returning the before twice as much power as has been calculated to produce Dam, with the potential for export to Gorges Three China’s to Grand Inga, Inga I and II were As a precursor Europe. to constitute constructed in 1972 and 1982, respectively, generating capacity. hydroelectric second largest Africa’s dams, lack of large As is the fate of many of the world’s operating at maintenance meant that by 2006 both were Plans for the massive an estimated 20% of total capacity. for in 2009 with proposals replaced Grand Inga Dam were a Grand Inga Cascades, once the potentially catastrophic It recognised. of Grand Inga were impacts environmental of new wetland habitat and the fi to date Efforts by new reservoirs… opportunities provided dams have to counter the ecosystem impacts of large of attention to met with limited success due to the lack the poor quality anticipating and avoiding such impacts, the diffi and uncertainty of predictions, and with all impacts, and the only partial implementation success of mitigation measures.’ on the Congo is the Grand Inga projects One of the largest the waters River (Showers 2009). The idea of diverting electricity generators was fi of the Congo through periodically in the 1920s and has been revived proposed second largest since that time. The Congo has the world’s fl signifi Figure 1.4. Main purposes of dams in Africa (data from FAO, Aquastat 2010a). Aquastat FAO, (data from in Africa of dams purposes 1.4. Main Figure 20 CHAPTER 1 | BACKGROUND Figure 1.5.ReductioninthesizeofLakeChadover theperiod1963to2007.Source: Musa the shores of reservoirs, infection canreach epidemic Nigeria indicate thatamongstcommunitiesresettled along disease (seealso Chapter4,thisvolume).Examples from providing favourable conditions fortransmissionofthe with artifi for thedisease,breed inslow-fl malaria. Snails,whichrepresent theintermediatehost to bethesecondmostdamaging economicallyafter of theseisschistosomiasis,aparasiticdiseaseconsidered vectors ofcertainimportantdiseases.Themostcommon reservoir provides more extensivehabitatsfortheaquatic with damconstructionarise,inparticular, astheupstream In tropical regions, problems ofpublichealthassociated small clupeids. by otherspeciesfavouringlacustrineconditions,suchas populations adaptedtorunningwaters,andreplacement fl is clearthatthetransformationofecosystemfrom ultimate stateadoptedbythefi ultimately decreases sharply. Itisdiffi fi and epiphyticalgaeincrease. However, thisexpansionof the sameway, algalgrazersdowell,asphytoplanktonic new conditions,andsupportsanincrease innumbers.In food formostofthegeneralistfeedersthatdowellin which are favoured bydrowned trees, becomesabasic of larvaeandnymphs lacustrine conditions.Forexample,thehighproliferation increase inthepopulationsofthosefi Soon aftertheclosure ofadam,there isasubstantial those observedinnorthtemperateman-madelakes. 1988) and,ingeneral,theimpactsare different from uviatile tolacustrineresults inthedisappearanceoffi sh populationsdoesnotlastlongandthefi cial lakesassociatedwithdamconstruction Povilla adusta owing orstagnantwater, sh community, butit shspeciesfavouring cult topredict the (Ephemeroptera), sh biomass sh sh over-abstraction anddiversion ofwaterfor agriculture, in and desertifi 5% ofitsvolume sincethe1960s,duetoboth drought vividly illustratedbyLakeChad, which hasshrunktoaround in otherpartsofAfrica.Thescale oftheproblem ismost agricultural uses,althoughthe problem isalsopresent by theover-abstraction ofwaterforurban,industrial and Africaare andsouthern bothheavily impacted Northern Water abstraction lower Volta (AkosomboDam,Ghana)(Petr1986). (Ryder 1978),andadeclineinoysterpopulationsthe fi has beenobserved,forexample,asadrasticdeclinein salinity (Jackson of nutrientsfl as areduction offreshwater infl reductions inestuarineandfreshwater fi Larger water-diversion projects inAfricahavealsoledto observed intemperateregions. and differ substantiallyfrom impactsofimpoundments societal impactsofalltheseeffects are veryimportant, they are inundatedandthenexposed.Theecological in large areas oflandfl This drop canamounttoseveralmetres, andcanresult room forincomingfl resulting from theopeningofsluice-gatestomake from impoundmentistheannualdrop inlevel(draw-down) Finally, apeculiarityoftheartifi (e.g., Ofoezie2002;OgbeideandUyigue2004). proportions, withupto80%ofthecommunityinfected sh populationsinthelowerNile(AswanDam,Egypt) cation caused byshortageofrainfalland the owing intodownstream areas andincreases et al ood watersduringtherainyseason. . 1988).Theresponse tosuchactions uctuating betweenperiodswhere cial tropical lakesresulting ow reduces theamount et al. 2008. s population, sh CHAPTER 1 | BACKGROUND 21 ow in oating aquatic plants, . 1993). Increasingly, sedimentation . 1993). Increasingly, sh species, fl et al ne sediments from one river to another, one river to another, ne sediments from shes; and © JOHANNES FÖRSTER Changes in the physical and chemical status of the Changes in the physical and chemical status the transfer of clear water such as through river water, which is likely to increase to a normally muddy river, algal growth; of fi Introduction leading to a loss of habitats, such as gravel spawning beds for fi fi of alien The spread and animal diseases and their vectors. the recipient river, with major implications for channel river, the recipient to a integrity and ecological functioning leading loss of biodiversity; frequent Flow reduction in the donor river and increased fl in the donor river and increased Flow reduction

Deforestation and sedimentation and Deforestation rapidly population has grown Since the 1950s, Africa’s 2.5% for the rate of around with an average growth period 1950-2005 (UN 2010). This demographic pressure such as around has accelerated rates of deforestation, of eastern Lakes the Great Africa. The case of Lake sedimentation increased is well known, where Tanganyika in changed species has resulted following deforestation composition and disruption of community interactions in (Cohen some areas from watershed deforestation, road building and other building and road deforestation, watershed from the head of the River Chari, a tributary of Lake Chad’. The the head of the River Chari, a tributary of and bids for the feasibility studies have been completed, opened in 2008. were project with water Generally speaking, the risks associated transfers can be summarized as follows: ● ● ● ● ow at shermen nally in 2008 . 2008) (Figure . 2008) (Figure et al . 2008). With more than . 2008). With more et al . 2008). The ‘Transaqua’ project envisions the project . 2008). The ‘Transaqua’ et al cient supplies. The obvious solution from a manager cient supplies. The obvious solution from construction of a ’broad navigable canal which, running construction of a ’broad along the eastern and northern of the Congo crest the waters of the extreme catchment, could intercept north eastern edges of the basin and, after a course of the Congo-Chad watershed about 2,400km would reach fl its entire in Central African territory and discharge particular for rice production (Musa particular for rice production Lake Chad around all wetlands in the region 1.5). Virtually up, and the of drying either dried up or on the verge are has been planned irrigation development of 213,400ha stopped at 33,824ha (Musa farmers, fi 20 million people, most of whom are dependent upon the lake and its and livestock breeders wetlands for their subsistence and livelihoods, surrounding being tabled is so serious that plans are the problem Water other rivers in the region. to transfer water from additional present here, transfers, such as proposed to species diversity (see to threats in relation problems transfers’, below). section ‘Water transfers Water Africa has left The uneven distribution of rainfall across have while others in water resources rich some regions insuffi water-rich point of view is to transfer water from or user’s water is limited. At present, catchments to those where In limited to southernhowever, Africa; this action is largely to divert the Ubangi River the 1960s, a plan was proposed the drying and dying Lake Chad. into the Chari to revitalize proposed water transfer schemes were Such inter-basin periods in the 1980s, 1990s and fi at different (Musa Deforestation of the Upper Guinean rainforest in western Ghana. Population growth across many parts of Africa has led to increased of deforestation levels and increased sediment loads in catchments. 22 CHAPTER 1 | BACKGROUND the eutrophication ofLakeVictoria (e.g.,Witte reviewed byNyenje of eutrophication across inlandsub-SaharanAfricaare issue September/October2008),andmanyexamples eutrophic (Water Research Commission,SouthAfrica; it isestimatedthat28%ofalllakesandreservoirs are quantities normallyrequired bytheecosystem.InAfrica, phosphorus) are released intofreshwaters inexcessof Eutrophication occurswhennutrients(mainlynitrogen and Eutrophication impacting Africa’s inlandwatersare discussedbelow. apparent (Dejoux1988).Themajorformsofwaterpollution and theirimpactonfreshwater communitiesisclearly types ofpollutionare rapidlyincreasing onthecontinent carried outinAfrica,despitethefactthatsources ofall Few detailedstudiesofwaterpollutionimpactshavebeen 1.2.2.2 Pollution particularly thoseofrock-dwelling communities. anthropogenic activitiesisimpactinglacustrinehabitats, many surfacewaters. a majorsource ofnutrients,leadingtoeutrophication of untreated intheenvironment, are increasingly becoming industries inurbanareas, whichare oftendischarged in sub-SaharanAfrica,wastewatersfrom sewageand and urbanareas. Indevelopingcountries,suchasthose nutrients causingeutrophication originatefrom agricultural (see Box1.1IsNileperch ascapegoat?).Mostofthe on LakeVictoria, Kampala,Uganda. Nile perchfi sh headsbeingsmokedatafi et al . (2010).Awellknownexampleis ©SARAHEMCC sh landingsite et al . 2008) aquatic environments where theNileperch cohabits Such aphenomenon wasnotobservedinthose other Sahelo-Sudan area, start toproliferate inLakeVictoria? Why didtheNileperch, whichoccursalloverthe ecosystem (Balirwa but alsomajorenvironmental changestothewholelake involving notonlytheimpactsofspeciesintroductions however, shownthatthereality ismore complex, as the‘Frankensteineffect‘. More recent studieshave, this newspecies.Suchaphenomenonisreferred to deal withthehighlyintensivepredation pressure from because theindigenousfi Nile perch wastoblameforthis‘ecologicalcatastrophe’, 1992). Theconclusionappeared obvious:theintroduced of endemiccichlidspeciesstrongly receded (Witte beginning ofthe1980s.Atsametime,populations into thelakein1960s,startedtoproliferate atthe of analienspecies.Thislarge predatory fi consequences onindigenousfaunaoftheintroduction Victoria isusuallyquotedasanexampleofthedisastrous The introduction ofNileperch ( Box 1.1IsNileperch ascapegoat? with afew, suchas mercury, showntoaccumulateup Heavy metalsare takenupbybothplantsandanimals, Heavy metals (see box1.1IsNileperch ascapegoat?). area ofLakeTanganyika andaround muchofLakeVictoria pollution are, forexample,wellknownintheBujumbura sub-Saharan Africa(Nyenje of wastewateristreated insewagetreatment plantsin anoxia.Lessthan30% demanding andcauses,inturn, cause primaryorganic pollution,whichishighlyoxygen- use. Sewageandothereffl a steepincrease inurban,industrialandagricultural land Population growth overthelastfewdecadeshasled to Domestic andindustrialorganicloads sulphide. generating harmfulgases,suchasmethaneandhydrogen of macrophytes, decompose,consumingoxygen and may result asdeadorganic matter, suchasthedie-off and animals.Inextreme cases,atotaldepletionofoxygen harmful secondarymetabolitesthatare toxictohumans organisms withintheaquaticcommunity, andmayproduce cyanobacteria candisruptvirtuallyallinteractionsbetween following deoxygenationofthewatercolumn.Blooms and ultimatelytheeradicationofspeciessuchasfi with cyanobacteria,large-scale bloomsofmacrophytes, increases inphytoplankton,replacement ofdiatoms The environmental effects ofeutrophication include et al . 2003). sh faunawasnotprepared to uents, richinorganic material, et al Lates niloticus . 2009).Suchformsof sh, introduced sh, ) toLake et al sh, . CHAPTER 1 | BACKGROUND 23 and sh fauna . 1995). In Lake cant impact on sh communities et al cant environmental environmental cant shes would probably shes would probably Stolothrissa tanganicae shery has been for pelagic sh community based on a predator sh community based on a predator was populations invertebrates sh and sheries shing in African lakes are reviewed by reviewed shing in African lakes are cant impact on the endemic fi cant impact The overall effect was summarised as a The overall effect in the local selective decline of large cant role y was found to have a signifi y was found to have species and an increase in clupeids (Coulter species and an increase Lates shing is often recognised as an important form of shing is often recognised of more than twenty years, and concluded that the effect that the effect and concluded twenty years, than of more on fi larvicides of the study in the Okavango 1997). In another low (Lévêque of deltamethrin for control the insecticide Delta, spraying fl the Tsetse non-target aquatic invertebrates, although subsequent although subsequent invertebrates, aquatic non-target for of likely recovery has found indications monitoring 2004). and Ramberg most species (Perkins 1.2.2.3 Impact of fi The impacts of fi in Pitcher and Hart (1995). a number of contributions Overfi of few documented examples are perturbation, but there following overexploitation. species completely disappearing in lake fi Impacts on species composition Craig commonly documented (e.g., more however, are, to have reported 1992). For example, demersal trawling is played a signifi and Ligtvoet (Goudswaard cichlid species in Lake Victoria 1988) (see below) and Malawi (Tweddle the main fi where Tanganyika, steady reduction species, the initial impact was a reported in all to a single community was reduced the 1991). Ultimately, pairing between predator-prey Lates stappersi. a stable fi change from system to an unstable community structured and prey with the visual sexual recognition system of many with the visual sexual recognition likely loss of cichlids, leading to hybridisation and anoxia limits genetic diversity (Spinney 2010). Water cichlids build where grounds the potential reproduction deep some species to move from nests, and has forced from the risk of predation water into the shallows where increased. is the Nile perch would in Lake Victoria of Nile perch The proliferation change in a deep and progressive from thus result to pollution. the ecological status of the lake, related of of the ecosystem led to the increase Transformation an abundant food by providing communities predator the shrimp biomass exploded as a (for example, source plankton populations). Even without of increased result indigenous Nile perch, predatory of the introduction endemic populations of cichlid fi of the signifi have declined as a result has most changes within the lake. Although the Nile perch likely had a signifi of the lake, the situation is not as clear cut as originally thought. This case study demonstrates why we need to be cautious when developing our conclusions about the species on native biodiversity. of introduced effects

cant cant et al. shed shed cant impact ies over a period sh fauna was overfi species occupy similar niches . 1992). sh (Amaize 2010). Considering sh (Amaize cation plants has resulted in an cation plants has resulted sh, environmental contamination, et al shing settlement where over 2,000 over where shing settlement Lates sh species without any signifi on their populations. For example, the same ecological ecological on their populations. For example, the same where imbalance was not observed in Lake Tanganyika, four autochtonous to the Nile perch. a number of major changes undergone has Lake Victoria since the beginning of the 20th century (Balirwa with other fi the indigenous fi 2003). Firstly, was a high there (see Section 1.2.1.3). Secondly, human population in the lake basin, in the local increase established. cotton and tea plantations were large where Development of the lake basin in this manner has led a number of pollutants, including in to increases fertilisers and insecticides as used in the plantations, of domestic and urban pollutants. loading and increased A lack of water purifi the lake, leading accumulation of pollutions throughout The main consequences of eutrophication. to widespread in water transparency, a reduction include eutrophication (lack of oxygen) in deeper water. anoxia and increased These two phenomena have had particularly signifi impacts on the endemic cichlid fauna. Increased water water impacts on the endemic cichlid fauna. Increased has interfered especially near the lake shore turbidity, sh ponds in Delta State (Nigeria), affecting an estimated (Nigeria), affecting sh ponds in Delta State families from different parts of the country make a living, different families from the cost is undoubtedly high. Pesticides of contamination in African lakes An important source use of pesticides to curb and rivers is the increasing for agricultural diseases of people and livestock, and of pesticides on the impact purposes. In Africa, however, communities is the functioning of aquatic ecosystems and in westernstill poorly documented. One long-term study of a weekly looked at the impact Africa has, however, application of insecticide against blackfl and damage to the fi and damage to the Oil pollution of oil, mainly in estuaries and lagoons, discharges Chronic to aquatic systems. An a major threat considered are demonstrating the serious impacts unfortunate example in July in inland waters was reported oil pollution can have about 6,000 may have destroyed 2010, when an oil spillage fi of fi USD100 million worth fi the massive loss of the food chain. Mercury discharge in freshwater systems systems in freshwater discharge Mercury chain. the food problem a particular mining, to gold linked is generally the in Africa, River system. Nevertheless, in the Congo low metals is still relatively of trace or heavy occurrence such as parts of the world, to some other when compared (Biney South America 24 CHAPTER 1 | BACKGROUND Lake Victoria FisheriesOrganization. Figure 1.6.Evolutionoffi Only a few specieshave beendeliberately introduced al to beamajorthreat tofreshwater biodiversity(Dudgeon The spread ofalieninvasive plantsandanimalsisrecognised 1.2.2.4 Invasivealienspecies longer term(Witte new fi catch compositionposethequestionofhowstable tonnes by2007.However, suchdramaticchanges in niloticus Oreochromis newly introduced species(particularly species declined,newfi (Fryer andIles1972)(Figure 1.6).Inthiscase,asthenative 1970s, wellbefore theNileperch wasestablished inthelake of theendemictilapiaspeciesbybeginning increased fi (nylon gillnetsandoutboard motors),combinedwith In thecaseofLakeVictoria, improved fi impacts ofnaturalenvironmental change. structure are apparent butare hard toseparatefrom the 1991). Insummary, theimpactsoffi through abalanceofcompetitiveinteractions(Coulter . 2005;Leprieur shery, basedonnon-native species, willbeoverthe shing effort, resulted inasevere drop incatches et al et al ). Thetotalcatchreached 0.9million . 2009;Samways andTaylor 2004). . 2007). sheries were established,targeting sh catchesinLakeVictoria (Kenya,Uganda,Tanzania). Source: FishStatPlus,FAO and shing oncommunity Lates niloticus shing technology shing and et et threat. Smallmouth bass( alien fi Cape province. InSouthAfrica(see Darwall Africa, where bassand trout were introduced intheWestern alien speciestoimprove sportfi 1995). Inthepast,however, fi newly created water-bodies suchasLakeKariba (Marshall replenish overexploited wildstocksortoboostfi The maingoalofdeliberateintroductions hasoftenbeento sustainable fi tilapias and bodies. Forinstance,innewlybuiltreservoirs, stockingby group forimproving fi the introduction oftilapiaspeciesasanimportant of theargument, thoseinvolvedinaquaculture encourage fi are reported tohavenegativeimpactson theindigenous fi niloticus body toanother. Forexample,Niletilapia, been speciestranslocationswithinAfricafrom onewater- Madagascar (Benstead into Africanfreshwaters untilnow, withtheexceptionof sh fauna(Canonico sheries throughout mostofinter-tropical Africa,where they shes introduced foranglingare aparticularly serious , hasbeenintroduced bothforaquaculture and Heterotis niloticus sheries (Moreau sheries et al sh production inmanyAfricanwater et al . 2005).However, ontheotherside Micropterus dolomieu . 2000).More often,there have shery offi et al has enabled development of hasenableddevelopmentof sheries, suchasinSouth . 1988). cers alsointroduced Oreochromis Oreochromis et al ), together sheriesin . 2009), CHAPTER 1 | BACKGROUND 25 © IUCN/DANIÈLE PERROT MAÎTRE nancing for better farming have only just begun to be considered (see www. have only just begun to be considered The impacts of climate change freshwaterbiodiversity.eu). detail in Chapter 8, Section 8.6.1 in more treated are Revolution and wetlands 1.2.3 The African Green of millions of African farmers, with hundreds of millions Tens living in subsistence conditions. Until of dependents, are to Africa’s donors sent only food aid in response recently, proposing deepening agricultural crisis. Now they are through agricultural production a new plan: increased Revolution. It will require Green African a homegrown four kinds of assistance: fi inputs; extension services to advise farmers on the new technologies; community nurseries to diversify production; laid out As previously and investments in infrastructure. for 2025 (UN-Water/Africa Vision in the Africa Water scale of 2003), this will translate into an unprecedented ), M. ), all and sh fauna . 2005), but Oncorhynchus et al Tilapia sparmanii Tilapia , have had major M. salmoides Lepomis macrochirus sh ( sh Salmo trutta sh often surviving in refuges of less than in refuges sh often surviving eld fed by Tiga Dam outside Kanoon, north-east Nigeria. Increased Dam production through an African agricultural eld fed by Tiga ) and bluegill sunfi and brown trout, trout, and brown with largemouth and spotted bass ( with largemouth punctulatus and banded tilapia ( North America, from in Africa, now dominate the fi further north from throughout more than 80% of the Olifants River system, more throughout with indigenous fi Rainbow trout, 1km in headwater streams. mykiss, impacts in cooler headwater streams. in more treated The impacts of invasive alien species are detail in Chapter 8, Section 8.2.3. 1.2.2.5 Climate change human water change and increasing climate Anthropogenic on available stress to place great widely expected use are Africa (Thieme across water resources biodiversity changes on freshwater of these the effects Irrigated tomato fi Such efforts will be greatlyby the aided Green will lead to unprecedented Revolution water sector. development within the provision of information on the distribution, ecology and conservation needs of freshwater species. 26 CHAPTER 1 | BACKGROUND wetlands are ofmajorimportanceinagriculturalsystems; management systems.Forexample,manyvalleybottom they are inextricablylinkedtocropping andlivestock ecosystem services,includingfood.Inmanyplaces people inAfricathrough theprovision ofnumerous role insustainingthelivelihoodsofmanymillions aesthetic qualities.Theseecosystemsplayanimportant immense biodiversityandhaveimportantrecreational and environmental quality, sustainlivelihoods,support Assessment 2005;UNEP2010).Wetlands maintain are valuableecosystems(MillenniumEcosystem It isbecomingincreasingly acceptedthatinlandwetlands 1.3 10, Section10.1. Africa’s watersectorare covered inmore detailinChapter waters, asprovided bythisstudy. Developmentplansfor needs offreshwater speciesthroughout Africansinland information onthedistribution,ecologyandconservation will begreatly aidedbyprovision ofreliable andaccessible ecosystems andtomitigatepotentialdamage.Suchefforts therefore, bemadetosafeguard theservicesoffreshwater development withinitswatersector. Everyeffort must, provided by wetlands, directly supportinglocal livelihoods. provided bywetlands,directly Boys holdingafi ecosystems inAfrica Functions andvaluesoffreshwater sh catch (Mormyrops spp.) in Bomane, Lower Aruwimi, D. R. Congo. Fish caught for food are a vital service avitalservice spp.)inBomane, LowerAruwimi,D.R.Congo. Fish caughtforfoodare sh catch(Mormyrops © KLAAS-DOUWE DJIKSTRA that makesthemvaluableinsocio-economicaswell range ofnaturalresources thatoccurwithinwetlands a large diversityoflife(Silverton and amultiplicityofhabitatsthatenablethemtosupport in theworld,characterizedbyhighprimaryproductivity Wetlands are amongstthemostproductive ecosystems 1.3.1 Wetland functions chemical components,andtheirsurrounding catchment. on theinteractionsbetweentheirphysical,biologicaland hence provide different ecosystemservices,depending people. Different wetlandsperformdifferent functionsand which bringbothphysicalandnon-physicalbenefi through theprovision ofarange‘ecosystemservices’, directly supportingandsustaininglivelihoods.Theydo this society. InAfrica,wetlandsplayaparticularlyvitalrole in of theecologicalfunctionstheyperformwithhuman The valueofwetlandsforpeoplearisesfrom theinteraction season andduringdroughts. a source ofarablelandorgrazingthroughout thedry key resource forcultivatorsandpastoralists,byproviding extended periods.Asaresult, thesewetlandsact as a into thedryseason,allowingcultivationofcrops for saturated duringthewetseason,ground retains water et al. 1999).Itisthewide t to ts CHAPTER 1 | BACKGROUND 27 © KEVIN SMITH lled with sand mined from the river bed by hand, loaded loaded bed by hand, the river mined from lled with sand on to the bank, and then transported in trucks to be used in in trucks to be used and then transported on to the bank, construction. Sand mining in the Sanaga River, Cameroon. The boats are The boats Cameroon. River, Sanaga mining in the Sand fi 28 CHAPTER 1 | BACKGROUND the concentration ofpollutants isreduced before the wetland, thepapyrus vegetationabsorbsnutrients, and Kampala. During thepassageofeffl effl the Nakivubopapyrusswamp receives semi-treated forms andremove nutrients from thewater. InUganda, and volatilisation)mayconvert heavymetalstoinsoluble (including nitrifi and biologicalprocesses thattakeplacewithinwetlands improving waterquality. Thelarge numberofchemical Some wetlandsplayanimportantrole inmaintainingor wetland functionsispresented inTable 1. some ofthephysicalbenefi and, hence,theconditionofwetland.Asummary and isdirectly related tothespecifi way inwhichtheydosoiscomplexandmultifunctional, and intangiblebenefi Wetlands inthiswayprovide awidevarietyoftangible purifi dry seasonriverfl of benefi recognized bycommunities,butincludeawidediversity use. Otherecosystemservicesare oftennotexplicitly materials; andwildplantsforfoodmedicinal fi ‘provisioning services’,suchas:domesticwatersupply; The physicalbenefi spatial scales(Lawton1999). evapotranspiration) across arangeoftemporaland (for example,nutrientcycling,condensationand an importantrole inmaintainingwetlandprocesses it isnowrecognized thatbiodiversity and thewetland’s locationwithinacatchment.However, specifi unique toanyparticularwetlandanddependentonsite- an ecosystem.Theexactnature ofthefunctionsis and abiotic(i.e.,soilwater)componentsthatcomprise through theinteractionofbiotic(i.e.,fl chemical andbiologicalprocesses thatoccur Ecological functionsare theresult ofthephysical, disadvantages. that overalltheadvantagesofwetlandsoutweighany do enhancehumanwellbeing,anditiswidelyaccepted al. reduce theprevalence ofinfectiousdiseases(Chipps ecosystems andassociatedbiodiversitywillgenerally are disturbed,whereas thepreservation ofintact carrying speciesmostoftenincrease whenwetlands It shouldbenoted,however, thatpopulationsofdisease one reason whysomanywetlandshavebeendrained. as mosquitoesandsomespeciesofmollusc–thisis to people,theydoalsoharbourdiseasevectorssuch 2005). Althoughwetlandsare generallyofgreat benefi ecological terms(MillenniumEcosystemAssessment sheries; livestockgrazing;cultivation;construction 2006;Keesing uent from thesewageworksof capitalcity cation, climateregulation andsedimentretention. c factors,includingclimate,geology, topography ts suchasfl cto, denitrifi cation, et al. ows, groundwater recharge, water ts derivedfrom wetlandsinclude ts tolarge numbersofpeople.The ood reduction, maintenanceof 2010).Manywetlandfunctions ts humansocietiesgainfrom cto, ammonifi cation, c ecologicalfunctions uent through the ora andfauna) per se , plays cation et t them a large numberdependonwetlandhabitatstosupport bird thatmigratebetweenEurope andAfricaeachyear, important formigratorybirds. Ofthe300speciesof stages oftheirlife-cycle.Wetlands are especially drought orfoodscarcity existelsewhere oratparticular them asrefuges, migratingtothemwhenconditionsof swamp tortoiseandlechweantelope),whilstothersuse animals livepermanentlyinwetlands(forexample,the Wetlands supportarichdiversityoffl also generatingincomeforlocalpeople. attractions, notonlyenrichingthelivesofvisitors,but as theOkavangoDeltainBotswanaare majortourist cohesiveness ofcommunities.Elsewhere, wetlands such Such ceremonies are importantinenhancingthesocial high waterandbackagainwhenthelevelsfall. the peoplemoveoutoffl year, importantculturalceremonies are conductedwhen hydrological dynamicsoftheBarotse fl Zambia hasdevelopedasaconsequenceofthe example, theculture oftheLozipeopleinwestern cognitive developmentandaestheticexperience.For functions. Theseare associatedwithspiritualenrichment, People alsogain the city(KansiimeandNalubega1999). water entersLakeVictoria, theprincipalwatersource for very often gounrecognised and are oftenunder-valued. accruing outside monetarysystems.Consequently, they Many ecosystem servicesare forms of‘publicgood’, depend. the conditionofnaturalenvironment onwhichthey this makesthemparticularly vulnerable tochangesin livelihood options,and often havelimitedalternative employment andculturalintegrity. Suchcommunities through provision offoodsupplies,medicines, income, critical contributionstotheirlivelihoodsandwelfare directly dependentonwetlandfunctionsthatprovide gained from them.InAfrica,manymillionsofpeopleare on socialperceptions oftheuseandbenefi The valuesattributedtowetlandsare largely dependent 1.3.2 Wetland values development andresource planningandmanagement. functions haveinthepastoftengoneunrecognised in people beingaware ofthem.Asaconsequence,wetland wetland contributionstohumanwelfare accruewithout far beyondtheboundariesofwetlanditself.Many wetlands andveryoftenthebenefi Many groups ofpeoplebenefi million birds eachyear(Roggeri1995). that theInnerDeltaofNigerhostsupto1.5 Africa eachsupportover400species,anditisestimated en route . TheOkavangoDeltaandKafueFlatsin non-physical oodplain duringperiodsof t from thepresence of benefi ora andfauna.Some ts are experienced ts from wetland odli. Each oodplain. ts tobe CHAPTER 1 | BACKGROUND 29 4HE NG LDSOF EITY NDFOR NLAND EOFAN ISTURE ION S ANDAREA THE EVERAND P AROUND MORE $ELTA LANDS WAS et al. . et al. et al et al (Zwarts -1 et al.

t human populations. 4HETOTALCATCHFROMINLANDWATERSIE LAKES RIVERSANDWET MILLIONTONNESIN&!/ 4HISCOMPARESTOMILL sheries and 48.4 million tonnes from fi marine capture tonnes from many people cannot afford in Africa, where However, aquaculture. sheries (2 the contribution of inland wild fi to practice aquaculture, than that of greater million tonnes) to the livelihoods of people is much CULTUREDlSHERIES TONNES 5.%0 #ATCHESFROMI subsistence in particular those from very often unreported, sheries are fi important to the rural poor in particular. of great sheries, which are fi 4HE(ADEJIA .GURUWETLANDSIN.IGERIACONTRIBUTETOTHERECHARG that live 1 million people aquifer used for water supply by approximately (Hollis in the region "ASEDONTHEINCLUSIVE2AMSARDElNITIONOFWETLANDS OVERHALF world’s population (i.e., more than 3 billion people) obtain their basic people) obtain their basic than 3 billion (i.e., more population world’s billion 3 wetlands. The remaining freshwater inland from water needs via cases is recharged which in some on groundwater, people depend wetlands. )NTHE+ILOMBEROmOODPLAINWETLANDIN4ANZANIA OF@POOR HOUSEHOLDSANDOF@BETTER OFFHOUSEHOLDSRELYONTHEWETLA DRINKINGWATER-C#ARTNEYANDVAN+OPPEN )NWESTERN!FRICA THESHALLOWGROUNDWATERANDELEVATEDSOILMO used valley wetlands are of inland of about 5 million hectares cultivation. extensively in crop .ON IRRIGATEDRICEGROWNONTHEmOODPLAINSOFTHE)NNER.IGER mUCTUATESBETWEEN AND TONNESPERYEAR WITHYIE THEORDEROF &ORMAT+GHA DIVERSITYOFORGANISMS INCLUDINGOVER SPECIESOFPLANTS THANSPECIESOFBIRDS ANDSPECIESOFlSH(OWARD +AFUEAND,UENA&LATS WETLANDSIN:AMBIA SUPPORTANOUTSTANDI 7ETLANDSAREIMPORTANTINTHEBIOGEOCHEMICALCYCLINGOFCARBON DIOXIDE METHANEANDHYDROGENSULPHIDE+AYRNALI 'LOBALLY WETLANDPEATDEPOSITSOCCUPYJUSTOFTHEWORLDSL BUTSTORE OFTHEPLANETSSOILCARBONPOOL,LOYDINPRE ,OCALPEOPLEUSETHEFRUITS SEEDS TUBERS ROOTSANDLEAVESOF 3EWAGEFROMOFTHERESIDENTSOFTHECITYOF+AMPALAAROUND PEOPLE ISDISCHARGEDINTOTHEKM§.AKIVUBOWETLAND the quality of water cantly improves of the wetland signifi presence main supply the city’s 3km from entering Lake Victoria, approximately INTAKE+ANSIIMEAND.ALUBEGA PLANTSFROMTHEWETLANDSSURROUNDING,AKE#HILWAIN-ALAWI )NTERNATIONAL7ATER-ANAGEMENT)NSTITUTE 4HEWETLANDSOFTHE+AFUEmOODPLAININ:AMBIASUPPORTlSHERIE AVERAGING TONNESPERYEAR-C#ARTNEY ,OCALPEOPLECOLLECTEIGHTPLANTSPECIESFROMTHE"UMBWISIDI FRESHWATERWETLANDIN4ANZANIATOTREATAILMENTSRANGINGFROMF to chest pains and coughs (Rebelo stomach disorders 'LOBALLY WETLANDSAREHIGHLYPRODUCTIVEAND BECAUSEHETEROGEN in a wide variety of ecological and soil conditions results in hydrology niches, they support immense biodiversity (Junk s s s s s s s s s s s 7ATERSUPPLY s 7ATERSUPPLY !GRICULTURE &ISHERIES 'ASREGULATION s 7ASTEWATERTREATMENT s &OODANDMEDICINE "IODIVERSITY s s s s s s s Groundwater recharge Groundwater Wetland functionWetland Storage of precipitation Use by society and runoff Examples Chemical cycling .UTRIENTTRANSFORMATION s Biomass production sh (e.g., wood, grass, fi etc.) Table 1.1. Examples of wetland functions that benefi that functions of wetland 1.1. Examples Table 30 CHAPTER 1 | BACKGROUND The OkavangoDeltainBotswanaisamajorglobaltouristattraction,generatingincomeforlocalcommunities. 2004; Emerton2005;SchuytAdekola (Barbier have beenmadeatboththemicro andmacro scales Attempts tovaluesomewetlandecosystemservices a year(UNEP 2006).In addition,the valueofwetland value interms of crops aloneisclose toUSD50million considerable, with estimatessuggestingthe economic value ofwetlandsintheZambezi Riverbasinisalso Zambia’s GDPin1990(Seyam estimated tobetheequivalent ofapproximately 5%of agriculture accounting forthemainshare) havebeen wetlands (withfi they provide. Forexample,thetotalusevalueofZambia’s values havebeenassignedtotheecosystemservices values extremely diffi functions, however, makesquantifi The site-specifi opportunity costsofmaintainingthemintact. wetland ecosystemsare generallyfargreater thanthe These havedemonstratedthatthereplacement costsfor et al. 1997;MitschandGosselink2000;Terer sh production andfl c nature anddiversityofwetland cult. Forsomewetlands,monetary et al. 2001).Theeconomic oodplain recession oodplain cation ofwetland et al. 2008). such traditional systems are increasingly underpressure sustainable use ofwetlands(Terer been established tosafeguard resources and ensure have political arrangementsandsettlement patterns in manycasesfi are ofteninharmony withhydrological regimes, and customs. Traditional resource managementstrategies whole, whichofteninforms traditional practicesand about wetlandresources andtheenvironment asa many placesthere isagreat dealoflocal knowledge the socialvaluesofwetlandsare alsoconsiderable.In determinant oftheirvalue.Beyondthepurely fi services provided bywetlandswithmoneyasthesole It isimpossibletoassessallofthefunctionsand provided byAfricanwetlands. that havebeenderivedforparticularecosystemservices 2 provides someadditionalexamplesofmonetaryvalues production valuedatoverUSD70millionannually. Table a year, whilethefl fi sheries inthebasinisestimatedtobeUSD80million oodplain grasslandssupportlivestock shing cycles,andpeoples’socio- et al. © WILLIAMDARWALL 2004).However, nancial, CHAPTER 1 | BACKGROUND 31 ts WETLANDSIN.IGERIAIS ON NTHElSHERIESOFTHE)NNER IN:AMBIAAREESTIMATEDTO ELY HOUSEHOLDSAND MEOFTHE7ESTERN#APE 3OUTH CATTLETHATGRAZEONTHE MILLIONPERYEAR3CHUYT WANAEACHYEAR GENERATINGAN COSYSTEMIN:AMBIAHASBEEN ANWETLANDIN#APE4OWN 3OUTH LUDINGMAIZE RICE SWEET EDTOGENERATEAGROSSINCOME FRICA WITHATOTALVALUEOF53$ HEYCATCHBETWEEN TO CANNOTAFFORDTOPRACTICE AAREVALUEDAT53$MILLIONPER et al . et al . 2009). This diversity and complexity of . 2009). This diversity and complexity of et al rewood. It has been shown, however, that the use that the use It has been shown, however, rewood. et al. and plants for traditional medicines, building materials and fi location markedly from differs of wetland resources socio-economic groups to location, as well as across (Rebelo complicates the development of wetland utilization greatly management policies. If the benefi wetland appropriate wetlands bring are to be used more wisely in the future, wisely in the future, to be used more wetlands bring are insight into the dynamics much greater is need for there of wetland functions and the links between wetlands and such understanding will the Only through human society. true value of wetlands be determined. ected ected &ISHERIESINTHE,AKE#HILWAWETLAND -ALAWI AREVALUEDAT53$ FROMWILDRESOURCESOF53$MILLIONPERYEAR3!2$# 4HETOTALAREAOFWETLANDSIN4ANZANIA HA ISESTIMAT .IGER$ELTAAmOODPLAINWETLAND $EPENDINGONmOODEXTENTT 4HELIVELIHOODSOFAPPROXIMATELY PEOPLEAREDEPENDENTO TONNESPERYEAR:WARTS ISESTIMATEDTOBEWORTH53$MILLION)NTHESAMEAREA mOODPLAINAREVALUEDATAPPROXIMATELY53$MILLION%MERTON 4HEESTIMATEDTOTALGRAZINGVALUEOFTHEHA-FULENIPERI URB !FRICA IS53$ ,ANNASAND4URPIE !GRICULTURALACTIVITIESINTHE(ADEJIA .GURUWETLANDSIN.IGERI YEAR3CHUYT )NCONTRASTTOTHEGLOBALlGURES IN!FRICA WHEREMANYPEOPLE /NTHE"AROTSEmOODPLAIN :AMBIA HAOFCULTIVATIONINC POTATOES SUGARCANE FRUITANDVEGETABLES SUPPORTSAPPROXIMAT of people sheries (2 million tonnes) to the livelihoods wild fi the contribution of inland aquaculture, ISMUCHGREATERTHANTHATOFCULTUREDlSHERIES TONNES 2EEDSANDPAPYRUSCOLLECTEDFROMTHE"AROTSEmOODPLAINWETLAND HAVEAVALUETOLOCALCOMMUNITIESOF53$ PERYEAR%MERT 4HEECONOMICVALUEOFGROUNDWATERRECHARGEINTHE(ADEJIA .GURU Africa, is estimated to be USD43.7 million (Turpie ESTIMATETOBE53$ PERYEAR!CHARYAAND"ARBIER 4HEANNUALWATERTREATMENTVALUEOFWETLANDSINTHE&YNBOS"IO 4HEECONOMICVALUEOFTHEWETLANDBIODIVERSITYOFTHE"AROTSEE ESTIMATEDTOBE53$ 3CHUYT !PPROXIMATELY TOURISTSVISITTHE/KAVANGO$ELTAIN"OTS of the primary tourist attractions in southernannual income of USD13 million. This makes it one Africa. 4HEREARE LINElSHINGPERMITSSOLDANNUALLYIN3OUTH! shing. for inland fi shing, an unknown number are for marine fi million. Although many of these are sh on inland rivers and fi exploitation of freshwater no data, recreational are Although there WETLANDSISKNOWNTOBEEXTENSIVE&!/ sheries, dry season s s s s s s s s s s s s s ji, Tana, Niger and Zambezi river catchments each ji, Tana, Fibre and fuel Fibre Fisheries Ecosystem ServiceEcosystem Provisioning: Agriculture Value Regulating: recharge Groundwater Pollution control and detoxifi cation and detoxifi Pollution control Supporting: Biodiversity Cultural: Recreational as population rises, people’s need for cash income need for cash income as population rises, people’s management institutions and contemporary increases (for example, formal government) ones based replace on custom. of wetland ecosystems is refl The high productivity in the high human populations that they can support. The wetlands of the Nile, Congo, and estuarine freshwater Rifi of thousands sustain the livelihoods of many hundreds of fi provision of people through grazing for cattle, water for domestic use and agriculture Table 1.2. Examples of values of African wetlands. of African of values 1.2. Examples Table Chapter 2. Assessment methodology

Darwall W.R.T and Smith K.S.¹

¹ IUCN Species Programme, 219c Huntingdon Road, Cambridge, UK

MAIN: Chutes Guillaume on the Kwango River, a tributary of the Kasai which forms the Angola and D.R.C. border © DENIS TWEDDLE RIGHT: Ledermanniella spp. are habitat specialists found in waterfalls, rapids and riffl es. They require well oxygenated and clean water and are therefore very sensitive to changes in river fl ows © J.-P. GHOGUE CONTENTS 2.1 Selection of priority taxa 34 2.1.1 Fishes 34 2.1.2 Molluscs 34 2.1.3 Odonates 34 2.1.4 Crabs 35 2.1.5 Aquatic plants 35 2.2 Regional approach 36 2.3 Data collation and quality control 37 2.4 Species mapping 38 2.5 Assessment of species threatened status 39 2.6 Nomenclature 41 34 CHAPTER 2 | METHODS defi For thepurposes ofthisassessment,freshwater fi fi access tofood,incomeandlivelihoods from smallscale al. consumption offi Africa is,however, the onlycontinentwhere theper-capita Africa(WorldFishthe majorriversofwestern Center2008)). alivingasfi (for example,165,000peopleearn communities andemploymentincomeformanymore 2006), providing essentialnutritionforthepoorest of nearly 25%oftheworld’s inlandcapture fi million tonnesperyear(FAO 2006).Thisaccountsfor estimated thatinAfrica,inlandfi for manymore (Coates1995;Dugan worldwide (FAO 2002)andfoodsecurityemployment the primarysource ofprotein fornearly1 billionpeople for poorpeople”(WorldFish Center2005).Theyprovide at aglobalscale,andare oftenreferred toasa“richfood Arguably, fi 2.1.1 Fishes ecosystems. indication oftheoverallstatusassociatedwetland conservation status,whencombined,willprovide auseful taxonomic groups, informationontheirdistributionsand levels andecologicalroles encompassedwithinthesefi nonetheless important.Giventhewiderangeoftrophic the othertaxamaybeindirect andpoorlyappreciated but income orasavaluablefoodsupply, benefi many peoplethroughout theregion, eitherasasource of Although fi and aquaticplants. molluscs; odonates(dragonfl information. Thetaxonomicgroups selectedwere: fi there wasthoughttobeareasonable levelofpre-existing Priority groups were selectedtoincludethosetaxaforwhich food websthatunderlieandsupportwetlandecosystems. selected torepresent arangeoftrophic levelswithinthe Therefore, anumberofprioritytaxonomicgroups were noticed. Clearly, itisnotpracticaltoassessallspecies. ecosystems, eveniftheyare neithercharismaticnor often essential tothemaintenanceofhealthyfunctioningwetland to conservethoseothercomponentsofthefoodweb we takeamore holisticapproach bycollatinginformation have received mostattention.Itis,however, importantthat of aquaticsystems,itisthewetlandbirds andfi charismatic groups, suchasmammalsandbirds. Inthecase to humansthrough direct consumption,orthemore often includingthosegroups thatprovide obviousbenefi have focusedonalimitedrangeoftaxonomicgroups, most In themajorityofcases,large-scale biodiversityassessments 2.1 Selectionofprioritytaxa sheries are likelytodecline(WorldFish Center2005). 2003).Asaconsequence,while populationsincrease, ned asthosespecies thatspendalloracriticalpart sh provide aclearbenefi shes formthemostimportantwetlandproduct sh isprojected todecline(Delgado ies anddamselfl sheries landnearly2.5 t tothelivelihoodsof et al. ts provided by 2010).Itis shermen on sey (FAO shery ies); crabs; crabs; ies); shes that that shes shes are shes; shes; ve ts et habitat selection ofadult dragonfl (order Odonata)are dependent onfreshwater habitats. The Larvae ofalmost allspeciesofdragonfl 2.1.3 Odonates species (679species)ontheIUCN RedListatthattime. assessed before this assessment, withjust14%ofknown these, onlyasmallnumberhadtheirconservationstatus undescribed gastropod species(Strong valid descriptionsexist,inadditiontoapossible4,000 mollusc species(Bogan2008;Strong and snails).There are justfewerthan5,000 freshwater (e.g.,clamsandsomemussels as foodorornaments species are ofhighcommercial importance tohumans 2004; Howard andCuffey 2006;Vaughan to otheranimals(seeStrayer through fi to theircontributionwaterqualityandnutrientcycling essential tothemaintenanceofwetlands,primarilydue a keyrole intheprovision ofecosystemservicesandare parasites anddiseases.Thisisunfortunate,astheyplay (as avector)inthetransmissionofhumanandlivestock in anegativelight,assomespeciesplaysignifi rarely attractingtheattentionofpopularmedia,unless not normallyconsidered asbeingcharismaticcreatures, Vaughan threatened groups ofanimals(Lydeard Freshwater molluscsare oneofthemostdiverseand 2.1.2 Molluscs using theIUCNRedListCategoriesandCriteria. assessed for7%(918species)offreshwater fi project in2005,theriskofglobalextinctionhadonlybeen included (Lévêque or about15,000speciesifbrackishwaterare are almost13,000freshwater fi confi of theirlifecycleinfresh waters.Thosespeciesentirely population. tothe use ofsmallmeshnets(mosquitonets)posesathreat locality inLakeMai-N'dombe(Leopold Hemichromis serasogaster ned tobrackishwatersare alsoassessed.There lter-feeding, algal-grazingandasafoodsource et al. © ANTONLAMBOJ 2004).Theyare mostlyunobtrusive,andare et al. 2008).Priortothestartofthis (EN), only known from the type (EN),onlyknownfromthetype et al. sh speciesintheworld, ies strongly depends on 1999;Vaughan II), D.R et al. ies anddamselfl et al. et al. 2008)forwhich . Congo.The . et al. 2008).Some sh species sh 2008).Of cant role cant 2004; et al. ies

CHAPTER 2 | METHODS 35

et ned et al. 2008). 2009). oating (a et al. et al. . 2008; Vié . 2008; Vié et al ts to human communities 2004; Cumberlidge 2004; Cumberlidge (LC), a widespread species throughout et al. © DENIS TWEDDLE uke) in Asia, Africa, and the Neotropics. The fact the Neotropics. uke) in Asia, Africa, and . 2009), and an intermediate host of paragonimiasis . 2009), and an intermediate . 2002; Abdallah . 2002; Abdallah ltration and nutrient recycling. An aquatic plant is defi ltration and nutrient recycling. 2008). Only 20 species of aquatic plants had been assessed2008). Only 20 species of aquatic plants had this assessment began. for the IUCN Red List before Freshwater crabs are vectors of the parasite that causes of the parasite vectors crabs are Freshwater in Africa (Cumberlidge (river blindness) onchocerciasis et al (lung fl is a food-bornethat paragonimiasis indicates zoonosis widely consumed by humans. crabs are that freshwater good indicators of water quality, crabs are Freshwater habitats and found in wide variety of aquatic as they are (Yeo pristine water conditions relatively require 9% (174 species) of than slightly more Only a small number, status assessed crabs, had their conservation freshwater began. this assessment before for the IUCN Red List 2.1.5 Aquatic plants the building blocks of wetland Aquatic plants are food, oxygen and habitats for many ecosystems, providing also a hugely important natural other species. They are benefi direct providing resource, highly aquatic plants are the world. Numerous across structuralvalued for their nutritious, medicinal, cultural, also key species in the They are or biological properties. of wetland ecosystem services, such as water provision fi Sudanonautes aubryi western Africa. al as a plant that is physiologically bound to water (a here plant whose photosynthetically or as a terrestrial hydrophyte) or fl active parts tolerate long periods submerged to Cook (1996) aquatic helophyte) (Cook 1996). According between 1 and 2% of the approximately plants represent to between300,000 species of vascular plants, equivalent 2,900 and 5,800 species (Chambers cycling role may be taken up by freshwater crabs, as the crabs, freshwater up by may be taken role cycling of most in the diet of detritus importance overwhelming high biomass indicates their abundance and species and (Dobson in African rivers key shredders that they are

et et al. © KEVIN SMITH ow, turbidity or ow, (LC), widespread in Africa (except . 2009), of which 120 species are recognised from Africa. Africa. from recognised . 2009), of which 120 species are the terrestrial vegetation type, and their larvae develop in the terrestrial to water with regards they play a critical role water where and aquatic habitat structure. nutrient cycling, quality, as often regarded predators, voracious The larvae are of insect pest species. A full array important in the control as and, within the group of ecological niches is represented to changes in water fl susceptible they are and Rowe 2009), they loss of aquatic vegetation (Trueman have been widely used as an indicator for wetland quality. even 5,680 extant described species. However, are There is well studied, it is believed that the though the group actual number is close to 7,000 species (Kalkman al guild was thought to be almost The detritus shredding African systems. most tropical completely absent from it is now thought that this important nutrient However, 2008). Fewer than 1% (31 species) of odonates had had their risk of extinction assessed using the IUCN Red List started. Categories and Criteria by the time this project 2.1.4 Crabs crabs (Cumberlidge 1,280 species of freshwater are There Acisoma panorpoides in dense rain forest), southern Europe, the Middle East, southern and the Indian Ocean islands. Asia, 36 CHAPTER 2 | METHODS Figure 2.1.Regionalassessmentboundariesbased onmajorriverbasins. at theglobalscale. number ofspecies;andv)thefamilyiswidelyrepresented wide rangeofecologicalnichesandcontainasubstantial iv) theselectedfamilieswould,whencombined,covera the relevant species;iii)thetaxonomyisrelatively stable; ii) there isareasonable levelofavailableinformationon contains arelatively large proportion ofaquatic species; families wasbasedonthefollowingcriteria:i)family within thesefamilywere assessed).Theselection of although notcomprehensively (i.e.,notallaquaticspecies Chapter 7),withspeciesfrom otherfamiliesalsoassessed, species from 21selectedplantfamilieswasassessed (see For thisproject, theconservationstatusofallaquatic plant of north-eastern Africa)highlightingthestatusand of north-eastern completed, areport waspublished(withtheexception (see Figure 2.1).Onceeachregional assessmentwas into sixregions, delineatedbymajorrivercatchments would needtobeassessedatonetime.Africawasdivided and iii)toensure onlyamanageablenumberofspecies engagement withregional scientistsanddecisionmakers; (as mightbelostattheglobalscale);ii)tofacilitatebetter i) toallowregional conservationprioritiestobeidentifi a regional approach wastakenforthefollowingreasons: To collateinformationonthefreshwater speciesofAfrica, 2.2 Regionalapproach ed CHAPTER 2 | METHODS 37 ned lists to sh working elds are based elds are cation schemes cation © KEVIN SMITH Assessment date cation-schemes. cation schemes allow for elds allowing the assessors to add general elds allowing the assessors to add general Biogeographic realm Biogeographic realm (CS) Assessor & evaluator names cation schemes using pre-defi cation schemes elds within SIS are summarized in Table 2.1; some summarized in Table elds within SIS are The western Africa regional review workshop (fi in 2006. group) held in Accra, Ghana a number region Following the training workshop, in each contracted to collate, and of participating experts were on each input within the SIS, all available information The required the priority taxonomic groups. species from data fi text fi free are habitat information, such as for species distributions, other fi whereas and ecology, preferences on classifi classifi against. Standard record or geographic other groups consistency in analysis across information on the classifi For more regions. www.iucnredlist. employed visit the IUCN Red List website org/technical-documents/classifi of species for the production sourced Spatial data were the from distribution maps (see Section 2.4). All species then assessed at the were selected taxonomic groups scale, using the IUCN Red List Categories and regional Criteria version 3.1 (IUCN 2001) and the Guidelines for cation Scheme cation occurrence (CS) occurrence measures (CS) measures Red List assessment rationale (text) 2010; Brooks 2010; Brooks . 2005; Darwall . 2005; et al et al. eld; CS = Classifi Species Information Service (SIS) database for the compilation of a elds within the Fields Criteria (CS) 2009; García 2009; García et al. ed by IUCN, the project partners, and through partners, and through ed by IUCN, the project 2011) and the data made freely available through available through the data made freely 2011) and 2009; Smith 2009; Smith Taxonomy Higher taxonomy Synonyms Common names Geographic range General information (text) Countries of PopulationHabitat and ecologyUse and trade General information (text) General information (text) Habitats (CS) General information (text) (CS) Population trend Utilisation (CS) System (CS) (CS) Harvest trends Movement patterns (CS) ThreatsConservation measures General information (text) Conservation Bibliography General information (text) (CS) Threats References Red List assessment Red List Category and nal pan-Africa workshop in 2009, at which point data from et al. were data sets regional Red List website. All the IUCN that is the pan-Africa data set in resulting then combined, and analysed here. presented in 2003, with the easternThe assessments began Africa funded project assessment acting as a pilot regional Affairs Ministry of Foreign separately by The Netherlands’ successful completion of the (DGIS). Following the were regions eastern Africa assessment, the remaining culminating in approach, a staggered assessed through Given the six- here. presented the continental assessment which the assessments have been year time period over that some earlier assessments completed, it is unavoidable those species may now be out of date, especially for made, however, were Efforts subject to ongoing threats. earlier assessments during the to update information from fi species and updated to provide merged were all regions the most current assessments at the continental scale. For information, please visit www.iucnredlist.org. et al. distribution of each taxonomic group and the conservation conservation and the group taxonomic of each distribution (see Darwall the region within priorities 2.3 Data collation and quality control 2.3 Data collation and quality and collated for all known Information was sourced (see Section species within the priority taxonomic groups Africa and beyond (as necessary) across 2.1). Experts from identifi were consultation with the relevant IUCN SSC Specialist Groups. IUCN SSC Specialist Groups. consultation with the relevant database, trained in use of the project These experts were application of the Species Information Service (SIS), (IUCN 2001), the IUCN Red List Categories and Criteria for digitally and Geographic Information Systems (GIS) mapping species distributions. Table 2.1. Compulsory data fi 2.1. Table species assessment. Text = text fi species assessment. Text 38 CHAPTER 2 | METHODS to individual river/lake sub-basins, as delineated by the to individual river/lakesub-basins, as delineated by the that couldonlybe mappedtocountryboundaries, mapped Species distributions were, withtheexception ofsomeplants 2.4 Speciesmapping Cairo inMarch 2009. were thenre-evaluated atafi data set.Allspecies(apartfrom thoseendemictoaregion) maps, were thencombinedtoproduce the pan-Africa All regional assessmentdata,includingspeciesdistribution List CategoriesandCriteriahadbeenappliedcorrectly. presented wasbothcompleteandcorrect; andii)theRed independent expertstoensure that:i)theinformation species assessmentwasevaluatedbyatleasttwo was thenreviewed atasecondworkshop,where each Version 3.0(IUCN2003) (seeSection2.5).Allinformation Application ofIUCNRedListCriteriaatRegionalLevels: (USGS EROS),asusedtomapandanalysespeciesdistributions. Figure 2.2.Level6riverbasinsasdelineatedbythe editedHYDRO1KElevationDerivativeDatabase nal review workshopheldin where aspecies isinferred/expected tooccuralthough its many speciesithas alsobeenpossibletoidentify sub-basins to currently containeachspeciesoffi used inmostcasestoidentifywhich sub-basinsare known has beenrecorded), expertopinion,andpublisheddatawere Point localities(thelatitudeandlongitude where thespecies appropriate management unitforinlandwaters. accepted thattheriver/lakebasinorcatchmentismost always extendthroughout ariversub-basin,itisgenerally as, eventhoughitisrecognised thatspeciesrangesmaynot selected asthespatialunitsformappingspeciesdistributions continental Africa(seeFigure 2.2).Riversub-basinswere data setidentifi before submissionforsubsequentanalysis.Theresulting database wasfi only pan-AfricaGISriverbasinlayeravailable.TheHYDRO1K At thetimeofdatacollationphase,thisrepresented the level 6(thehighestresolution available)usingGISsoftware. HYDRO1K ElevationDerivativeDatabase(USGSEROS)at ed 7,079individualsub-basinsinmainland rst editedtoeliminatemanysmallerrors sh, odonate or crab. For sh, odonate orcrab.For CHAPTER 2 | METHODS 39

To ensure ensure To . appended RG . RG lters were applied in this lters were . © KEVIN SMITH ve criteria with quantitative thresholds ecting biological indicators of populations lters” to exclude certain species from the assessment certain species from lters” to exclude Species having less than 5% of their global distribution not assessed, and; within Africa were

For an explanation of the full range of categories, and theFor an explanation of the full range of categories, under eachcriteria that must be met for a species to qualify documentation: to the following please refer Category, 3.1, Version The IUCN Red List Categories and Criteria: http://www.iucnredlist.org/ which can be downloaded from technical-documents/categories-and-criteria. is the application assessment process Part of the regional of “fi The following fi appropriate. where case: 1) that this regional approach was consistent and comparable was consistent and approach that this regional for the Guidelines assessments, regional to other IUCN Red List Criteria at Regional Levels:Application of IUCN Categories followed. Red List 3.0 (IUCN 2003) were Version 2.3. shown in Figure level are at the regional is considered “Critically Endangered” A species assessed as in the wild. high risk of extinction to be facing an extremely be to is considered as “Endangered” A species assessed of extinction in the wild. A species facing a very high risk to be facing a high is considered assessed as “Vulnerable” the wild. All taxa listed as Critically risk of extinction in described as are or Vulnerable Endangered Endangered, threatened distinguish between the three To “threatened”. fi are categories, there 2.3), refl (Table with extinction threatened “Regional” as opposed identify which assessments are To assessments have to “Global”, the regional for example, anin superscript to their Red List Category; species would be written EN Endangered for any species endemic to continental Africa represent the Africa represent continental endemic to species for any with those species risk, but for extinction global species’ status Africa the Red List extending beyond distributions within part of the population to that relates assigned only Assessments” termed “Regional are Africa; these rmed rmed species 2008). This new et al. rmed. These “inferred” sub- These “inferred” rmed. 2008). The IUCN Red List of nal pan-Africa review workshop, held in Cairo, Egypt March 2009. et al. (IUCN 2001). Red List assessments completed

2006, Mace 2003; De Grammont and Cuarón 2006; Rodrigues 2003; De Grammont and Cuarón 2006; Assessment of species threatened Assessment of species threatened status presence has not yet been confi not yet been has presence with confi to those connected often are basins determined distributions were Inferred locality records. scale knowledge, course a combination of expert through For many information. and unpublished records, distribution are distribution maps and mollusc species, of the plant localities basins, as digitized point based on inferred entirely not available. Finally, information were or detailed distribution to mapped were plant species many of the widespread detailed the absence of any more country boundaries in distribution information. is based on species spatial here All analysis presented sub-basins delineated by HYDRO1Kdata mapped to the the described above. However, (edited by IUCN), as that will be available for downloadindividual species maps have the IUCN Red List website (www.iucnredlist.org) from data to a new global hydrographic since been transferred (Lehner called HydroSHEDS standard data layer, which delineates 18,689 individual sub-basins which delineates 18,689 individual data layer, accurate and contains fewer for continental Africa, is more elevation data of than HYDRO1K, being derived from errors a higher resolution. Participants from the fi 2.5 widely accepted The Red List Categories and Criteria are available foras the most objective and authoritative system (Lamoreux assessing the risk of a species becoming extinct et al. et al. Threatened Species™ is the world’s most comprehensive comprehensive most Species™ is the world’s Threatened on the global conservation status of information source to help inform plant and animal species, and is widely use conservation priority setting. The risk of extinction for each species was assessed IUCN Red List Categories and Criteria: to the according 3.1 Version 40 CHAPTER 2 | METHODS Table 2.3.Summaryofthefi ubro aueidvdas<5 <250 extinction inthewildtobe: <50 Indicating theprobability of E. QuantitativeAnalysis Number ofmature individuals Either: D. Very smallorrestricted population (b) (aii) OR AND atleast2ofthefollowing: B2. B1. B. GeographicrangeintheformofeitherB1(extentoccurrence) AND/ORB2(area ofoccupancy) A4. A3. A2. A1. C2. (up toamax.of100yearsinfuture) (ai) A. Populationreduction ubro aueidvdas<20<250<10,000 <2,500 <250 C1. AND eitherC1orC2: Number ofmature individuals C. Smallpopulationsizeanddecline Use anyoftheCriteria (a) (c) (b) Extreme fl uctuations inthenumberofmature individuals. Numberofmature Ae focpny(O)<1 m <50k² <2,000km² <500km² <10km² Area ofoccupancy(AOO) Acontinuingdecline Anestimatedcontinuing %individualsinone xeto curne(O)<10k² ,0 m <20,000km² <5,000km² <100km² Extent ofoccurrence (EOO) may notbeunderstood the timeperiodmustincludebothpastandfuture, andwhere thecausesofreduction maynothaveceased An observed,estimated,inferred, projected orsuspectedpopulationreduction (uptoamaximumof100years)where (e) underAl. Population reduction projected orsuspectedtobemetinthefuture (uptoamaximumof100years)basedon(b) have ceased Population reduction observed,estimated,inferred, orsuspectedinthepastwhere thecausesofreduction maynot clearly reversible Population reduction observed,estimated,inferred, orsuspectedinthepastwhere thecausesofreduction are decline ofatleast: Severely fragmented, Extreme fl uctuations inanyof: Continuingdeclineinanyof: subpopulation = subpopulation: individuals ineach ubro oain =1 Number oflocations or subpopulations; of habitat; OR (iv) (e) (d) (c) (b) (a) effects ofintroduced taxa,hybridization,pathogens,pollutants,competitorsorparasites. adeclineinarea ofoccupancy(AOO),extentoccurrence (EOO)and/orhabitatquality direct observation numberoflocationsorsubpopulations; A2, A3&A4 actualorpotentiallevelsofexploitation anindexofabundanceappropriate tothetaxon maynotbeunderstood AND AND Declinesmeasured overthelongerof10yearsor3generations (iv) understood A–E ve Criteria(A–E)usedtodeterminetheCategory ofthreat foraspecies. OR OR (a)and/or(b): number ofmature individuals. A1 maynotbereversible, basedon(a)to(e)underAl. (i) OR (i) extentofoccurrence; rtclyEdnee nagrdVulnerable Endangered Critically Endangered AND > 25% in3yearsor1 extent ofoccurrence; 50%in10yearsor (100 yearsmax.) 3 generations haveceased,basedonandspecifyinganyofthefollowing: generation 010 510 100% 95–100% 90–100% OR > > 0<20<1,000 <250 < 50 80% 90% maynotbereversible, basedon(a)to(e)underAl. Restricted area ofoccupancy (ii) (v) (ii) area ofoccupancy; number ofmature individuals. area ofoccupancy; > 20% in5yearsor2 20%in20yearsor (100 yearsmax.) 5 generations generations > > 50% 70% < 5 (iii) (iii) area, extentand/orquality numberoflocations AND /OR D1. D2. 10% in10yearsor3 > <1,000 Typically: AOO 10%in100years of locations < 20km² generations > > < 30% 50% 10 or < number 5 OR

CHAPTER 2 | METHODS 41 © KEVIN SMITH sh Specialist Group, respectively. For plants, where For plants, where respectively. sh Specialist Group, A taxonomist at work at the South African Institute for Aquatic Biodiversity (SAIAB). single taxonomy for molluscs and crabs, and we therefore we therefore single taxonomy for molluscs and crabs, and by the IUCN SSC recommended follow the standards Crab and and the Freshwater Mollusc Specialist Group Crayﬁ Checklist of Selected Plant we follow the World appropriate, Kew (WCSP Families hosted by the Royal Botanic Gardens, also followed, specialist lists are 2010), but other more Ferns (Hassler and Swale such as the Checklist of World on the taxonomic standards information 2010). For more of the IUCN Red List, visit www.iucnredlist.org/technical- documents/information-sources-and-quality#standards. cation eld, and in many cases iteld, and in many cation generally follows the World Odonata cation generally follows the World nd a universally agreed taxonomic hierarchy. taxonomic hierarchy. nd a universally agreed cult to ﬁ Species arriving in Africa post 1500 were treated as treated Species arriving in Africa post 1500 were not assessed, but their being “non-native” and were possible. mapped where distributions were

Taxonomic schemes are constantly changing as results constantly changing as results schemes are Taxonomic in particular with the introduction ongoing studies, from is made available. Taxonomy of molecular techniques, are fi also a somewhat controversial is diffi 2.6 Nomenclature List maintained at the University of Puget Sound (Schorr no widely accepted is currently and Paulson 2010). There 2) presented are Species summaries and distribution maps for all species assessed on the DVD that accompanies the An example output is given in the paper copy of this report. Appendix. In this case, the taxonomy followed is that adopted possible, employs by the IUCN Red List which, where existing published world checklists. Fish classifi generally follows the online Catalog of Fishes maintained at the California Academy of Sciences (Eschmeyer 2010). Odonate classifi Figure 2.3. IUCN Red List Categories at the regional level. regional at the List Categories Red 2.3. IUCN Figure Chapter 3. The status and distribution of freshwater fi shes

Snoeks, J.¹, Harrison, I.J.² and Stiassny, M.L.J.³

1 Zoology Department, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080 Tervuren, Belgium and Laboratory of Animal Diversity and Systematics, Katholieke Universiteit Leuven, Charles Deberiotstraat 32 B-3000 Leuven, Belgium 2 Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA 3 Department of Ichthyology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA 3.1 Overview of the ichthyofauna of Africa 44 3.1.1 A short introduction to the ichthyofaunal regions 44 3.2 Conservation status 49 3.3 Patterns of overall species richness 49 3.3.1 All fi sh species 49 3.3.2 Threatened species 55 3.3.3 Restricted Range species 60 3.3.4 Data Defi cient species 63 3.3.5 Extinct species 65 3.4 Major threats to species 66 3.5 Research actions required 70 3.6 Conservation recommendations 71 Species in the spotlight – The Congo blind barb: Mbanza Ngungu’s albino cave fi sh 74 Species in the spotlight – Tilapia in eastern Africa – a friend and foe 76 Species in the spotlight – Forest remnants in western Africa – vanishing islands of sylvan fi shes 79 Species in the spotlight – A unique species fl ock in Lake Tana – the Labeobarbus complex 82 Species in the spotlight – The Twee River redfi n – a Critically Endangered minnow from South Africa 85 Species in the spotlight – Cauldrons for fi sh biodiversity: western Africa’s crater lakes 87 44 CHAPTER 3 | FISH crater lakes(see some notableexamplesintheEastAfricanGreat Lakes,the levels ofendemismare highformanypartsofAfrica,with rivers oftherainforestsandcentralAfrica.The ofwestern and thediversityofsympatricmormyridsfoundinsmall Lake Tana (seealso other examples,suchasthespeciesfl Verheyen Lake Victoria hasbeenheavilydebated(Stager fauna, theevolutionaryimpactofapossiblycompletelydry Victoria about14,700yearsagohadalarge infl et al lineage diversifi (Cichlidae) inLakeVictoria thatwentthrough amajor best-known caseisthe500ormore speciesofcichlids adaptive radiation,ifnoteven‘explosivespeciation’.The species groups thathaveundergone extensiverecent 35-54 millionyearsago.Africaalsohasrepresentative of theextantgeneraare younger, originatingintheEocene, years (LeCointre andLeGuyader2001),althoughmembers representatives. Fossillungfi the mostancientgroup ofbonyfi African speciesinthegenus (Lundberg (e.g., thebichirs,Polypteridae;lungfi Africa hasseveralarchaic andphylogeneticallyisolated taxa connection totheselandmassesaspartofGondwana. affi unique features compared toothercontinents,itshares Lévêque 1997).WhiletheAfricanichthyofaunashowsmany some populations,orextinctionforothers(Roberts1975; resulted ingeographicisolationfollowedbyspeciation for long historyofcomplexclimaticandgeologicaleventsthat Africa harboursawell-diversifi 3.1 further over thelasttwo decades,as a result ofseveral increase inknowledge,andthisknowledge hasexpanded (Boden [CLOFFA] (Daget waters, the inventory allthefi areas andpoorlydefi catalogue isamajorsource ofinformationforpoorlyknown most authoritativeaccountof1,425species.Evennow, this of theAfricanfi Museum inLondon.HisimportantfourvolumesCatalogue Belgian ichthyologist,thenworkingintheNaturalHistory the existinginformationwassetbyGeorge Boulenger, a is more thanacenturyold.Afi The scientifi the riversofcentralAfrica. ]fiÔ jg\Z`\jÕ nities withbothSouthAmericaandAsia,asaresult ofits . 2003). While it is clear that the desiccation of Lake . 2003).WhileitisclearthatthedesiccationofLake j_Y`f[`m\ij`kp1n\jk\ie8]i`ZXËjZiXk\icXb\j Africa Overview oftheichthyofauna et al fZbèCXb\KXeXÆk_\ et al c studyofAfrican fresh andbrackishwaterfi et al . 2004).CLOFFA represented aconsiderable Check-List of Freshwater Fishes of Africa Check-List ofFreshwater . 2004).Lundberg cation about100,000years ago(Verheyen shes (Boulenger1909-1916)provided the . 2000).Lungfi et al shes occurringinAfricancontinental

ned taxa.Inamore recent effort to Jg\Z`\jèk_\Jgfkc`^_k . 1984-1991)listed2,908species Jg\Z`\jèk_\jgfkc`^_k shes datebacksome410million ed fi Protopterus rst majorstepincompiling shes, whichincludefour sh fauna,resulting from a shes thatstillhaveliving et al CXY\fYXiYlj ock of se, Protopteridae) shes, . (2000)alsoprovide , are membersof Labeobarbus uence onthis – :Xlc[ifej – et al Zfdgc\o 8le`hl\ . 2004; . 2004; shes ) and ) and in )

and phylogeneticallyisolatedgroupoffi Tanganyika. withinthePolypteridaefamily, Bichirsare anarchaic throughouttheCongocatchmentandLake widespread Polypterus endlicheriicongicus ● ● ● ● ● ● ● ● ● ● Africa (Balian of thePalaearctic realm than withtheremaining partof continent hasa closeraffi (Doadrio 1994).Inbiogeographic terms,thispartofthe poor inspeciesnumbers,with dominanceofCyprinidae from otherregions ofcontinentalAfrica.Itisrelatively The Maghreb province inthenorthofAfricaisquitedistinct ichthyofaunal provinces (Figure 3.1): 1974); andMatthes(1964).Roberts(1975)recognised ten 1921, 1933);Nichols,(1928);Blanc(1954);Poll(1957, who basedhisworkonBoulenger(1905);Pellegrin(1911, synthesiswasformulatedbyRoberts(1975), a modern dates backmore thanacentury. However, thebasisfor Attempts tosubdivideAfricaintoichthyofaunalprovinces 3.1.1 Ashortintroduction totheichthyofaunalregions region, andtheCongolianAngolanriversystems. from theAfricancontinent,especiallyfrom theGreat Lakes likely thatseveralhundred speciesare stilltobedescribed endemic, asare abouthalfofthefamilies.Inaddition,itis not restricted tothespecieslevel;majorityofgeneraare almost allofthemendemictotherealm. Thisendemismis partoftheArabianPeninsula)isabout3,200, the southern Africa,butincludingMadagascarand excluding northern the EthiopianorAfrotropical Zoologicalrealm (thatis,Africa For example,currently inFishBase,thenumberoffi fi total numberofscientifi in FishBase(Froese andPauly, 2010),highlightingthatthe museum collectionsfrom around theworld,are synthesized below). Manyoftheseandotherstudies,aswelldatafrom et al Africa;andStiassny Skelton (1993)publishedforsouthern et al regional accounts.Lévêque shes hasrisendramaticallysinceCLOFFA wascompiled.

Southern (including CapeofGoodHope) Southern Quanza Zambezi East Coast Zaire (Congo)(includinglakesKivuandTanganyika) Lower Guinea Upper Guinea Nilo-Sudan Abyssinian (Ethiopian)Highlands Maghreb . (2007a,b)publishedfortheLowerGuineanregion (see . (2003a,b) published accounts for western Africa; Africa; . (2003a,b)publishedaccountsforwestern et al . 2008b; Lévêque cally knownAfricanfreshwater nity withthe Mediterraneanpart et al (LC), a subspecies of bichir (LC),asubspeciesofbichir . (1990,1992)andPaugy et al shes. . 2008). © ULISCHLIEWEN shes in in shes CHAPTER 3 | FISH 45 gure. gure. The (2007a). (1) (2007a). et al. Stiassny ed from 2007a,b), more than half of which are endemic. half of which are than 2007a,b), more et al. (haplochromines) should be added to this ﬁ (haplochromines) complete, existing inventory of the Nile system is far from especially in the poorly documented Sudanese part. In addition, the endemic cichlid fauna of Lake Albert contains still to be described (Snoeks pers. many species that are obs.). of the Nilo-Sudan and the border River forms The Cross While the river is considered the Lower Guinea province. its fauna includes to be part of the Lower Guinea province, (Stiassny and Hopkins 2007). elements of both provinces and brackish included 577 fresh The Lower Guinea province of its ichthyofauna review water species at the time of a recent (Stiassny River southwards the Cross from the area spans The region to just north of the Congo Basin. The Shiloango (Chiloango), as in Cabinda (Angola), is regarded with its lower reaches some However, basin of this area. the southernmost large a mixed fauna occur in smaller coastal basins with probably the Shiloango and the Congo. between the region Jg\Z`\j

; see fZb è CXb\ KXeX 2003a,b), including uents) and the Niger, uents) and the Niger, ), Albert and Turkana ), Albert and Turkana et al. Labeobarbus Zfdgc\o 8 le`hl\ jg\Z`\j Õ – ned (Lévêque 1997). In a recent review of the review ned (Lévêque 1997). In a recent CXY\fYXiYlj . 2009). However, several additional species that are that are several additional species . 2009). However, shes of western Africa (Paugy è k_\ jgfkc`^_k Æ k_\ Maghreb, (2) Nilo-Sudan, (3) Abyssinian Highlands, (4) Upper Guinea, (5) Lower Guinea, (6) Congo (Zaire), (7) Congo (Zaire), Guinea, (6) Lower Guinea, (5) (4) Upper Highlands, (3) Abyssinian Nilo-Sudan, (2) Maghreb, Coast, (10) Southern Zambezi, (9) East Quanza, (8) Good Hope). (including Cape of Figure 3.1. The major ichthyofaunal provinces of continental Africa, modifi Africa, continental of provinces ichthyofaunal major 3.1. The Figure the Upper Guinea and the Nilo-Sudan provinces, but the Upper Guinea and the Nilo-Sudan provinces, and brackish water excluding the Nile system, 584 fresh of the Nile Basin fauna is long listed. A review species were but counts of the River Nile (excluding the region overdue, include 128 species (Witte and Edward) of Lakes Victoria et al ( endemic to Lakes Tana The Nilo-Sudan province is the largest, spanning the total spanning is the largest, The Nilo-Sudan province Senegal to Mozambique. It width of the continent from includes two major river systems, the Nile (but excluding and their affl Edward and lakes Victoria African coastal basins, and the endorheic Lake many West spanning the coastal Chad system. It excludes a region Guinea to basins of the so-called Guinean ridge, from the western Guinea part of Ivory Coast. This is the Upper (see below), for which the exact boundaries are province not well defi fi 46 CHAPTER 3 | FISH Mekong have greater fi Mekong have greater The confl basin. royauxi caughtspecimenofthecatfi A freshly ichthyofaunal terms,LakeKivubelongstotheEastCoast and Tanganyika andtheMalagarasisystem.However, in includes partoftheRiftValley region, includinglakesKivu fi (Revenga andKura2003).Ascurrently estimated,its basin inAfricaandgloballyissecondonlytotheAmazon The CongoBasinhasthelargest catchmentarea ofany sh faunaincludesaround 1,250species.Intheeastit © JOHNFRIEL ) (LC)thatlivesinlargerapidstheCongoRiver uence of theInkisiRiverandlowersection oftheCongoRiver, D.R.Congo. OnlytheAmazonandperhaps sh species richness. sh ( sh Euchilichthys ©ROBERT SCHELLY its non-cichlidspecies(Snoeks2000;DeVos cichlids are endemictothelake,asare more than60%of ichthyofauna. More than95%ofitsapproximately 200 oldest large lakeinAfricawhichisrefl distinct andnoteworthysectionoftheCongoBasin.Itis province (Snoeks this system.However, thelake’s basinandtheUpper Shire (Skelton 1994).Geographically, LakeMalawiisalsopartof borderthe southern isdelimitatedbySt.LuciaBasin the Zambeziichthyofaunalprovince.part, Ontheeastern system andtheLimpopoare themajorcomponentsof The Cunene,theOkavangoBasin,large Zambezi several endemics. 109 freshwater fi known ichthyofaunalprovinces inAfrica.Poll(1967) listed is diffi north oftheCunene.Anestimatenumberspecies of thecoastalbasinsinAngola,southCongoand The Quanzaprovince isrelatively small,includingmost Congo speciesare endemic. the LakeTanganyika endemics, anestimated75%ofthe of mixedorigin(DeVos Its majoraffl cult togive,asthisregion isoneoftheleastwell- uent, theMalagarasicontainsanichthyofauna sh speciesfrom thisregion, including et al . 1997).LakeTanganyika isaquite et al . 2001).Evenwhendisregarding ected initsdistinctive et al . 2001). CHAPTER 3 | FISH 47 ect past . 1997). All ned mainly et al shing pressure. ned. The Cape province includes includes province The Cape ned. ock, comprising more than 700 endemic than more ock, comprising shing community. This iconic species is generally shing community. . 1997). While these provinces mostly refl . 1997). While these provinces the Orange system and all basins south of it. However, south of it. However, system and all basins the Orange Forty- Zambezi elements. also includes typical the Orange 36 of Cape province, occur in the well-known two species 2001). endemic (Skelton which are is situated to province The East Coast ichthyofaunal poor is relatively and province the north of the Zambezi by savanna. Skelton (1994) in species and dominated various river systems, with the from 125 species recorded 60%. These systems include all an endemicity of about system of the Zambezi to the Tana coastal basins north species richness and in northern Kenya. In this region as many of these underestimated, endemism may well be other main component of The underexplored. rivers are of various origins, includes a series of lakes the province and its Victoria Kyoga, George, Rukwa, Kivu, Edward, smaller satellite lakes (Snoeks numerous of a regional these lakes, except Rukwa, have elements super species fl cichlids, and many undescribed species of haplochromine species. was provinces Roberts’ (1975) work on ichthyofaunal (1983) and complemented continued by Greenwood 1997; Snoeks by others (e.g., Skelton 1994; Lévêque et al drainage patterns defi and current and are endemism. The border between the Cape and Zambezi and Zambezi the Cape between border The endemism. not well defi is provinces sh families. (LC), a species popular among the sport fi sh families, this is less evidently the Hydrocynus vittatus Hydrocynus sh, harbour a unique fauna dominated by some 800 or more 800 or more harbour a unique fauna dominated by some many more endemic cichlids (Snoeks, 2004), and probably The remaining undescribed species (see section 3.3.1). well known, is relatively part of the Zambezi province characteristically with species numbers in this region north to south (Skelton 2001). from decreasing includes many of the typically While the Zambezi province sub-Saharan African fi which case for the southern ichthyofaunal province, or Cape depauperate in families but high in species is relatively common and abundant with a wide distribution across Africa, but is locally depleted by heavy fi common and abundant with a wide distribution across Africa, but is locally depleted by © MELANIE L.J. STIASSNY The tigerfi © TRACY FARRELL The Chobe River, a major tributary The of the Zambezi. The Chobe River, province is relativelyZambezi ichthyofaunal well known and sub-Saharan African fi contains many typical 48 CHAPTER 3 | FISH Figure 3.2.Amapofthefreshwater ecoregions ofAfrica,from Thieme the region, the communities ofaquaticspecies present, characteristics, including thehydrological features of were defi renaming someofthe ecoregions) (Figure 3.2).Ecoregions Thysville cavesintotheLower Congoecoregion and this, recognising only 78ecoregions (incorporatingthe offshore islands). Abell on continentalAfrica(exclusive ofMadagascarand Thieme Figure 1.2). Africa, basedon‘freshwater ecoregions’ (See Chapter1, different systemfordefi zones oftenexist.Thieme boundaries are notalwaysstraightforward, andtransition on acharacteristiccombinationofendemictaxa,their et al ned bya combination ofphysicalandbiological . (2005)recognised 79freshwater ecoregions ning thebiogeographicregions of et al et al . (2008)subsequentlyrevised . (2005)developedaslightly García Africa (e.g.,Darwall in theassessments ofthestatusfreshwater speciesin been adoptedbyIUCNforthe freshwater fi describing speciesdistributions bysub-catchmentshas partial sub-catchments.Forthese reasons, themethodof complete catchmentsorsub-catchments, ratherthan for freshwater ecosystems are usuallyimplementedfor about aregion, conservationplanningandmanagement biological, ecological,andbiogeographicinformation While theecoregion approach provides veryuseful basins thatare componentsofquitedifferent catchments. may includepartialcatchments,oraggregate sub- exactly matchedtorivercatchments;insomecasesthey Consequently, boundariesofecoregions are notalways and associatedecologicalevolutionaryprocesses. et al . 2010a; Brooks et al. (2005) et al . 2005,2009;Smith et al . 2011). shes included shes et al . 2009; CHAPTER 3 | FISH 49 shes) sh shes recorded from from shes recorded shes of Africa according to shes of Africa according shes), with 198 species. shes), with 198 currently sh (at least 243 species are sh species sh ed in Nothobranchiidae and Poeciliidae), and the 3.3 Patterns of overall species richness 3.3.1 All fi of the numbers of Lévêque (1997) undertook a review and brackish water fi fresh Although the taxonomy (i.e., families, genera, species). study provides numbers will have changed since then, his 76 He recorded a useful overview of taxonomic diversity. fauna dominated by families in Africa, with the freshwater typically are ostariophysans (1,159 species), many of which by only represented several families are riverine; however, form the a few species. The Cyprinidae (475 species) of ostariophysans, and Characiformes proportion greatest by the Alestidae (109 species) and also well represented are Distichodontidae (90 species). The Siluriformes (catfi species of Mochokidae (176 species), include numerous and Clariidae (74 species). (98 species), Claroteidae Cichlidae is Among the non-ostariophysan groups, with at least 870 species the most species rich family, represented (1997); most of these are to Lévêque according by species endemic to the lakes of eastern Africa. Other numbers of species include the former families with large Cyprinodontidae, killifi classifi Mormyridae (elephantfi The geographic distribution of species shows some of very high richness as well as areas of distinctive areas 3.4). Not low richness, or even absence of species (Figure no fi are in this study there surprisingly, some of the driest parts of Africa, for example, much of the Sahara, parts of Ethiopia and Somalia, the Kalahari Desert parts of Namibia. In contrast, areas of Botswana, and large Figure 3.3. The proportion (%) of freshwater fi of freshwater (%) proportion 3.3. The Figure in Red List Category IUCN regional in each species continental Africa. mainland ed as sh ., 2007a,b; species endemic shes. Three Number of et al gures gures represent ed species) are ‘Data ed species) are sh species assessed sh species ed as ‘Least Concern’ed ., 2011). Number of species cant threats. Six hundred Six hundred cant threats. cient species are greatest greatest are cient species et al cient information about their . 2009; Stiassny shes face signifi species cient species. Most of the threatened ed as threatened – representing 21.8% of all – representing ed as threatened et al cient’ (DD), with insuffi Critically Endangered Critically Endangered 148Endangered 354Vulnerable Near Threatened cient Data Defi 148 117Least Concern 353 speciesTotal 117 75 510 1629 73 2836 509 1585 2788 IUCN Red List Category Extinct 3 3 shes in other areas (see below). Such undeveloped areas undeveloped areas (see below). Such shes in other areas Table 3.1. The number of African freshwater fi The number of African freshwater 3.1. Table species in each IUCN Red List Category. and nineteen species (with an additional 16 sub-species) and nineteen species (with an additional classifi are if one discounts assessed species, or 26.6% of all species the Data Defi classifi category, in the lowest threatened are Tweddle Tweddle of the classifi Over 500 species (18% Defi distribution to assess whether they ecology or taxonomy, the conclusion or not. This underscores threatened are surveying and that a considerable amount of additional to provide is required monitoring of African freshwaters accurate assessment of the conservation status a more in parts of of species in these ecosystems, particularly lakes of eastern Africa central Africa and the Rift Valley numbers of Data Defi where such before (see section 3.3.4 below). Nevertheless, even it is possible surveying and monitoring is implemented, that many to say that all existing evidence indicates fi freshwater species), with another (57.2% of all threatened ‘Vulnerable’ and species listed as ‘Endangered,’ 23.9% of threatened These fi 18.9% as ‘Critically Endangered’. 3.1), further highlighting numbers of species (Table large fi to African freshwater the severity of threats as ‘Extinct’, although this is probably reported species are of the true numbers (see section 3.3.5). an underestimate Of the 2,836 African freshwater fi African freshwater Of the 2,836 3.2 Conservation status 3.2 Conservation not Africa (that is, of mainland continental at the scale as ‘Not Applicable’ the four species ranked including classifi half (57.4% are (NA)), over be partly explained 3.3). This may 3.1, Figure (see Table sparsely populated of Africa that are areas by the large agricultural, industrial, or little is relatively there and where to threat a severe present currently urban development that fi Lower Guinea, parts of the Congo Basin, include large of southern Africa (Stiassny and regions 50 CHAPTER 3 | FISH number ofspeciesperriver/lakesub-catchment. Figure 3.4.Thedistributionoffreshwater fi mouth-brooding haplochromine species. Anestimated 67 of speciespresent are cichlids, mainlyrepresented by are not assessedfortheIUCNRed List.Themajority 3.3.3). However, many are notyetformallydescribedso species offi (Thieme connects thetwolakes,comprise theMalawiecoregion infl (358 specieshavebeenassessed).LakeMalawiandits assessments havebeencompletedfortheIUCNRedList province) hasthegreatest numberofspecies forwhich the large lakes,LakeMalawi(intheZambesiichthyofaunal the fauna),andmainchannelofCongoRiver. Of of theRiftValley Africa (where ofeastern cichlidsdominate of greatest speciesrichnessincludesomeofthelarge lakes uents, LakeMalombe andtheUpperShire Riverthat et al shes, mostofwhichare endemic(seesection . 2005).Thisregion includes anestimated800 sh speciesacross mainlandcontinentalAfrica.Speciesrichness= affl species, withatotal of54non-cichlidspeciesfound inthe of thesub-catchments around the lake are alsorichin present inthelake(Agnese andTeugels 2001).Several generalist species, 2004), appearingtohaveoriginated from awidespread of thegenus nine and12speciesofmostly deep-water large catfi microcephalum (mpasa), whichisEndangered, andatrout-like sardella cyprinids, includingasardine-like pelagic (Snoeks 2004).Thelakehassomeeconomicallyimportant 10 thatare yettobeformallydescribed)are notcichlids of thesespecies(includingtwointroduced speciesand uent rivers(Snoeks 2004). (usipa)(LC),asalmon-like Bathyclarias (sanjika)(VU)(Thieme Clarias gariepinus are endemic tothelake(Snoeks Opsaridium microlepis (LCRG),thatisalso et al . 2005).Between Engraulicypris shes O.

CHAPTER 3 | FISH 51 ocks of sh Stolothrissa . 2005). Thieme et al shes including endemic shes including endemic (LC) and shery in the lake (Thieme uents. (LC), a species of elephant fi (LC), a species of elephant sh species have been assessed for the . 2011). This number is certainly an uent drainages at the northern tip of the . 2011). For example, recent surveys in the . 2011). For example, recent (2005) note there may be more than 600 may be more (2005) note there Limnothrissa miodon et al (LC)) that are prey to several other species prey (LC)) that are et al et al. © JOHN FRIEL . 2005). Affl . (2005) note that the lake also has species fl . (2005) note that the lake also has species shes (Claroteidae and Mochokidae), snooks (Latidae) shes (Claroteidae Snoeks 1994; Snoeks 2000; Thieme et al catfi also supports and spiny eels (Mastacembelidae). The lake a unique community of pelagic fi clupeids ( tanganicae fi and which support an off-shore et al lower section lake support up to 43 species, while the the lake holds of the Malagarasi on the eastern of shore have been the highest number of species (71 species assessed) for any of the affl in Lake Victoria Estimates for the total number of species variable, although are (part of the East Coast province) Thieme of Malebo lower section of the Congo River downstream Mormyrops anguilloides Mormyrops cichlids, and several endemic species. Most of these are or Possibly Extinct (see section Critically Endangered are 3.3.2 and 3.3.5). Many of the endemic species of cichlids gone extinct since the 1980s (Harrison thought to have are and Stiassny 1999). The sub-catchments adjacent to Lake 21 and 51 species that have been hold between Victoria numbers found assessed for the Red List, with the greatest in the Nzoia drainage to the north-east of the lake. than 858 fi More Congo Basin (i.e., that part of the Congo ichthyological lakes Tanganyika) of the Rift Valley exclusive province (Stiassny poorly are many of these regions and underestimate, number is at all; the recorded or not explored explored year described each species are as more increasing (Stiassny widespread It is a member of the in Sub-Saharan Africa. one of the most species-rich families in Mormyridae family, Africa.

(EN) is from Algeria and Tunisia, Tunisia, (EN) is from Algeria and . (2005). The lake has high levels of endemism, . (2005). The lake has high levels of endemism, © FRANK DOUWES et al shes, 287 of which had been formally described from shes, 287 of which had been formally described from shes, most of which are endemic and from the family © ANTON LAMBOJ The Lake Tanganyika Basin (part of the Congo Basin Basin (part The Lake Tanganyika harbours an estimated 470 species ichthyofaunal province) of fi by reported the lake itself at the time these numbers were Thieme 300 of Around especially for cichlids (see section 3.3.3). cichlids, including species (64%) are the 470 recorded as well as species-rich lineages of substrate-brooding and cichlids (Coulter 1991; De Vos mouth-brooding Lake Malawi ecoregion includes an estimated 800 species of fi Cichlidae. Haplochromis desfontainii Haplochromis where it is found in warm, freshwater springs. It is a member the most speciose in Africa. of the Cichlidae family, 52 CHAPTER 3 | FISH The Malebo Pool, one of the most species-rich areas currently knownthroughouttheCongocatchment. currently The MaleboPool,oneofthemostspecies-richareas An excellent caseinpoint istheLéfi other partsof the basinasinventoriesare undertaken. and similarobservations are being madethroughout 2008; Monsembulapers.comm.; Schliewenpers.comm.), previously documented (Thieme example, foundthemtoharbour manymore speciesthan region suchasthe Inkisi, Nsele,andMpozohave,for Recent surveysofsmallriverbasins inthelowerCongo compared tothesmaller, more inaccessible,tributaries. has occurred inthelarger channelsoftheCongoBasin species islikelyanartifactofmore intensivesamplingthat biodiversity assessments.Thisapparent distributionof some sub-catchmentshadnospeciesrecorded inthese have fewerspecies;than30inmanycases,and smaller tributariesdistributedthroughout theCongoBasin region itselfhas316assessedfi more than250assessedspecies,whiletheMaleboPool greatest speciesnumbers,withseveralsectionshaving Congo, betweenBoyomaFallsandMaleboPool,hasthe Uele, andSangharivers.Themiddlesectionofthe River, aswellsectionsoftheLualaba,Kasai,Ubangi/ than 150speciesreported formostreaches oftheCongo have particularlyhighnumbersofspecies,withmore The maincoursesofthemajorriversCongoBasin 2011). than 10newspeciesinthelastfi documented there, includingtheidentifi Pool havemore thandoubledthenumberofspecies sh species.However, the et al ve years (Stiassny years ve . 2008;Wamuini ni River, from which cation ofmore et al et al . . widespread throughout theLowerCongoRiverbasin. widespread intheOdzala NationalPark,D.R.Congo,is quite captured This species of freshwater pufferfi This speciesoffreshwater of centralAfricacontributetothehighlevelsspecies and climaticstabilityoftheforested, moisttropical regions The equatoriallocation,large sizeandtherelative longevity harbour 140species(Ibala-Zamba2010). a thorough explorationofitslowerreaches, itwasfoundto virtually nospecieswere knownuntilrecently when,after © JOHNFRIEL sh, Tetraodon miurus ©ROBERT SCHELLY

(LC), CHAPTER 3 | FISH 53 ow sh . 1997). et al sh fauna of sh. Lungfi Pronothobranchius 2003a,b). It is probably 2003a,b). It is probably (Helfmann (LC) is an air breather (and (LC) is an air breather et al. sh (e.g., ows more or less from west to or less from ows more . 2007a,b). (LC), the African lungfi et al P. aethiopicus P. © TIMO MORITZ shes distributed through westernshes distributed through Africa; (2003a), there are 584 species of fresh and 584 species of fresh are (2003a), there (NT)) that have drought resistant eggs (Laleye resistant (NT)) that have drought Protopterus annectens et al. sh, are adapted to survive of drought by burrowing to periods the bottom of mud in drying pools and aestivating there for up to eight months. (Stiassny the region much of western Africa has been well studied, Similarly, and early especially over the latter part of the 20th century taxonomic revisions of production with the 21st century, and faunal guides (e.g., Paugy after southern areas Africa. large one of the better-known ecoregions, Africa includes 17 freshwater Western the western the Nilo-Sudan distributed through parts of to According and the Upper Guinea province. province Paugy brackish water fi component) were 521 of these species (the freshwater Entsua-Mensah assessed for the Red List (Laleye and 2009). While several river basins in western Africa have relatively is less uniformly rich in high species numbers, the region of central Africa. species than the Lower Guinea province which fl The Niger River, part of western Africa, and is Africa’s a large east across has a patchy species density pattern. longest river, third Those parts of western richest in species Africa that are and woodland tend to be the coastal basins, moist forests those that have fewer species are savanna, whereas drier and rivers conditions are found in the Sahel, where the exception of the Niger) or may fl smaller (with are often Fishes found in these drier regions only seasonally. For example, the show adaptations to periods of drought. lungfi Protopterus annectens Protopterus fi of a guide to the freshwater the production must take lungfuls of air occasionally in order to survive) must take lungfuls of air occasionally in order mud at the bottom of drying pools into and can burrow aestivating, usually for up to seven or and survive there, eight months; this may be extended experimentally to up to four years in also species of killifi are There kiyawensis and Entsua-Mensah 2009). sh . 2010; . 2011). et al et al . 2006; Thieme cation (Thieme et al . 2009; Markert et al owing to the Congo basin. The Dja owing . 2007a,b). The most species-rich © TIMO MORITZ et al 2010). et al. 2005; Brummett . 2008). In addition, the region has a complex mosaic . 2008). In addition, the region River headwaters were once captured of Lower by rivers Guinea, possibly the reason two regions why the share some ichthyological fauna. Rapids on the Dja River fl Rapids on the Dja River More than 550 species have been reported from Lower Lower from have been reported than 550 species More Guinea (Stiassny drainages are the lower part of the Sanaga (Cameroon), lower part of the Sanaga (Cameroon), the drainages are the Ogowe (Gabon), the upper Ngounie (Gabon/Republic of Congo), and the lower Kouilou systems (Republic of (and assessed) Congo), each having over 100 recorded species. Most sub-catchments of the Lower Guinean 50 and 100 assessed species. have between province short relatively These include several coastal rivers that are rich in fi disproportionately (i.e., 60km or less) and are et al often ecoregions, of habitats, contributing to 19 freshwater barriers between the habitats with distinct hydrographic of these factors (for example, waterfalls and rapids); all high species diversifi appear to promote et al. their small size. Lower Guinea has been to species relative a focus for ichthyological survey over the last 20 years. extensive taxonomic and revisionary This has promoted work, including the description of many new species and Although the species richness observed in the Congo Although the species richness observed lakes exceeds that Basin and the east African Rift Valley several are observed in any other part of Africa, there high species richness that have a relatively other regions noticeable This is particularly areas. distributed over large ichthyological over almost all of the Lower Guinean parts of western Africa (covering the and large province western and the Upper Guinea part of the Nilo-Sudan, ichthyological provinces). is adjacent to the Congo The Lower Guinea province some fauna, perhaps as share Basin, and the two regions of the headwaters of a consequence of historic capture Congo Basin rivers by Lower Guinean rivers (for example, headwaters by the Nyong, Ntem, Ivindo) of the Dja capture 1966; Stiassny (Thys van den Audenaerde richness in this area (Kamdem Toham (Kamdem Toham richness in this area Stiassny 54 CHAPTER 3 | FISH one oftheshortestlivedkillifi withlungfi typically foundinseasonalpans,whichtheyoftenshare The turquoisekillifi compared Africa, LowerGuinea, theCongo, to western Other partsof Africahavelowerspecies numbers lake margins. ofseasonalfl patterns Chad. Manyofthespeciesin the region are adaptedto recorded), both ofwhichare majoraffl Yedseram andlowerChaririverbasins(with72species Lake Chad(with69species)andtheregion coveringthe muted (seeabove).Anexceptiontothisistheendorheic River, are situatedintheSahel,where speciesrichnessis Africa, whichincludethemiddlereaches oftheNiger partsoftheNilo-SudanprovinceThe northern inwestern species (Thieme of LakeVolta) andhascontributedtothedeclineofsome ecology oftheregion (mostnoticeablybytheformation Akosombo andKpongdamshassignifi lake inriverineandmarginal habitats.Constructionofthe probably includesspeciesfoundindrainagesclosetothe itself; however, thisnumbermaybemisleading,sinceit assessments, 105fi Entsua-Mensah 2009).According torecent biodiversity Lake Volta) hasbetween 160and185species(Laleye Nigeria.TheVoltasouth-western ecoregion (including Mali), andsomecoastalcatchmentsfrom IvoryCoastto Niger andInnerDeltaecoregions (inGuineaand Guinea,SierraLeone,andLiberia),theUpper southern those intheUpperGuineaprovince (coveringpartsof Africahavemoreof western than70species,particularly section 3.3.2).Severalcatchmentsandsub-catchments Lagos Lagoon,includingsixthreatened species(see assessed intheOgunRiverbasinregion ofthe (see section3.3.2).To thewest,107specieshavebeen also someofthemostheavilyimpactedareas inAfrica been assessed).Thedeltaandsurrounding areas are found intheNigerDeltaecoregion (152specieshave The greatestAfricaare numbersofspeciesinwestern et al sh, sh speciesare present inthe lake Nothobranchius furzeri . 2005;andseesection3.3.5). ooding in thelakeandaround the shes (fourtofi cantly affected the ve months). ve uents ofLake (LC),fromtheBahiniNationalParkinMozambique.Thisbeautifulkillifi © SAIAB/ROGERBILLS apparently eggs,andare shes. Theylaydrought-resistant 39 inthe RuvuandRufi recorded andassessedfor theTana RiverbasininKenya, noteworthy exceptions tothis.Twenty ninespecies are lakes andaffl have lowspeciesrichness.Lake Victoria anditssatellite partoftheNilo-Sudan province,Highlands, andtheeastern Most oftheEastCoastprovince justsouthoftheEthiopian below thedam. apparently extirpatedfrom theformerparts oftheirrange Africa; seeabove)(seesection3.3.2),withmanyspecies of theNilesystem(justaswithLakeVolta inwestern dam hasbeendetrimentaltothefreshwater fi this elevatednumberofspecies,theoverallimpact Aswan Dam,havebeenrecorded andassessed. Despite Nile. Fortysevenspeciesfrom LakeNasser, formedbythe partofEthiopia)thatdraintotheWhite the westernmost and inthewetlandsaround GambelaNationalPark(in found upstream from KhartoumintheBlueNilesystem, species (where 40to50specieshavebeenassessed)are recorded intheNileBasin.Thegreatest numbersof There are someexceptionstothelowspeciesnumbers Sudan. swamps insouthern all oftheUpperNileecoregions), includingthevastSudd basin withinSudan(coveringsomeoftheLowerNileand numbers isthelimitedexplorationoflarge partsofthe Nevertheless, partofthereason forlowrecorded species and are unabletosupportlarge numbersofspecies. affl the ariditywithinmuchofNile’s catchmentarea, where than fi much oftheEthiopianHighlandsprovince, havefewer the UpperandLowerNileecoregions, andthroughout of itslength.Manyaffl fewer than30speciesare recorded andassessedformost River isthelongestinworld(RevengaandKura2003), and theRiftValley lakes discussedabove.WhiletheNile uents tendtobesmall,andmanyfl ve speciesrecorded. Thisisprobably arefl uent rivers(discussedabove) are themost uents ofthemainchannel,inboth ji river basinsinTanzania, and25 o intermittently ow sh is sh s fauna sh ection of ection CHAPTER 3 | FISH 55 sh species shes are totally absent shes are , together with around 40 Oreochromis placidus Oreochromis and (Figure 3.5) is largely focused in a band that runs along the in a band that runs along focused 3.5) is largely (Figure coast of western Guinea province Africa and the Lower Zambezi/ the throughout to D. R. Congo, Senegal from basins in the northernOkavango part of southern Africa, of the Rift Valley the river basins and lakes of through eastern basins Africa, also including some of the coastal of easterneastern Africa, and some basins in the and a few pockets of are southern parts of South Africa. There in central Africa, in species along the Uele River threatened in north-eastern Africa, and in the of Lake Tana the region Northwest Africa freshwater Atlantic and Mediterranean of northern region Africa in the Maghreb ecoregions of most throughout species The absence of threatened of northern of the Hornthe arid areas Africa, and parts of Africa (comprising Somalia and eastern Ethiopia), as well some parts of southern Africa (especially Namibia and Botswana) is unsurprising; fi (see above). Some other many of these regions from number of sub-catchments may have only a very small species) but in these cases species (e.g., fewer than three 3.3.2 Threatened species Threatened 3.3.2 fi freshwater of threatened The distribution ows from Lake Amaramba and forms the largest tributary of © SAIAB/ROGER BILLS ecting a shery. ect the intensity of Barbus atkinsoni, Labeo cylindricus . 2009). et al sh, sustain an important local fi the Ruvuma River. The Lugenda River Valley’s rich wildlife has led to development of the area rich wildlife has led to development as a destination for ecotourism. The Lugenda River Valley’s the Ruvuma River. Within the river itself, species such as other species of fi The Lugenda River is located in northern Mozambique, where it fl in the Ruvuma River on the border between Tanzania and Tanzania between border on the Ruvuma River in the basins east coast most of the Otherwise, Mozambique. contrast a striking than 20 species, providing have fewer on of Lower Guinea rich coastal basins with the species refl and probably of the continent, the other side and the well-documented of limited survey combination by eastern aridity experienced episodes of Africa. Southern and Africa encompasses the Quanza, Zambezi, 22 freshwater it includes provinces; Cape ichthyofaunal habitat types. The and some very diverse ecoregions general pattern richness is one of declining species most of the sub- the west and south. For example, towards Africa have fewer than 10 species. catchments of South is found in parts of the Zambezi Highest species richness than 50 species Lake Kariba (more from basin upstream and 80 species in many sub-catchments, recorded are the lake itself) and immediately upriver from present are the Okavango in the lowest parts of the basin; in parts of basin; the Buzi basin; some higher parts of the Limpopo and some and Save basins; the Incomati-Pongola system; some of smaller coastal basins in Mozambique. However, in part, to refl these densities appear, (Tweddle collection efforts 56 CHAPTER 3 | FISH richness =numberofspeciesperriver/lakesub-catchment. Figure 3.5.Thedistributionofthreatened freshwater fi resistant eggs. resistant periods ofdroughtbyhavingdrought- Algeria. Manykillifi (Cyprinodontidae family)endemicto aphanius, isaspeciesofkillifi Aphanius saourensis © HEIKOKAERST sh speciessurvive (CR), theSahara sh sh speciesacross mainlandcontinentalAfrica.Species CHAPTER 3 | FISH 57 native , (EN) Haplochromis shes from northern shes from . 2010b). et al Pseudophoxinus punicus (EN) and this decline. There are two species of fi are this decline. There Endangered. considered Africa that are desfontainii by groundwater threatened and Algeria, are to Tunisia which extraction, dams, water pollution and drought, (García the area widely affect species, Most of the Nile Basin lacks globally threatened with the exception of the upper part of the basin (adjacent High and in the vicinity of Lake Tana. to Lake Victoria), species (44% to 75% of the of threatened proportions and several found in Lake Victoria species assessed) are of the adjoining sub-catchments. Between 26% and 43% Lake of in the region threatened of assessed species are . 2010b). et al sh species that are threatened within each sub-catchment across across sub-catchment each within threatened that are sh species ) are the main reasons for the main reasons ) are ), is present in the Sahara freshwater Sahara freshwater in the ), is present Gambusia holbrooki sh ( sh Aphanius saourensi between 44% and 75% of the species are threatened; this threatened; between 44% and 75% of the species are and is the case for some sub-catchments in the Maghreb in in the Etosha and the Karstveld Sink Holes ecoregions 3.6). Namibia (Figure species, the Sahara aphanius One Critically Endangered ( is endemic to the Oued Saoura This species ecoregion. several parts of the from Basin, but has disappeared to a single population found basin and is now restricted near Mazzer in the Sahara desert (García pollution extraction for agriculture, Excessive groundwater of and proliferation and introduction wetlands, of remaining mosquitofi Figure 3.6. The proportion of freshwater fi freshwater of proportion 3.6. The Figure that are sub-catchment per river/lake species of = proportion richness Species Africa. continental mainland threatened. 58 CHAPTER 3 | FISH ranges. restricted endemic tothelake,andmanyhaveextremely species collectivelyknownas‘mbuna’.Theseare A LakeMalawicichlidfromthegroupofrock-dwelling document evidenceof current orimminent threats toa extinction. Given themore recent requirement toalso of offspring, whichmakethemparticularly vulnerableto such astheirhighlyrestricted rangesandlownumbers the ecologicalcharacteristics of manycichlidspecies, A numberoftheseassessments are, however, basedon species (28%ofthetotal assessedforthislake). Lake Malawi,where there are 105recorded threatened The highestnumberofthreatened speciesoccursin total speciesrichnessisstillrelatively high(26%ormore). draining totheKasai,ratioofthreatened speciestothe of theSanghaRiver, andtributariesoftheKwangoRiver on DataDefi assess threats tospeciesfoundthere (see section3.3.4 central Africaare alsoinsuffi in manypartsofthisregion. However, severalpartsof may bepartlyattributedtothelackofhumandevelopment of centralAfrica,includingthespeciesrichCongoBasin, The lownumbersofthreatened speciesrecorded formuch of theseextirpations. and theconstructionofAswanDamwasamajorcause Africa) were previously foundintheNileBasinEgypt, freshwater fi African (2010b) notethatatleast80%ofthe24northern Africanpartsoftheirrange.Garcíain thenorthern species inthebasinhaveundergone seriousdeclines the rest oftheNileBasin,itisimportanttonotethatseveral globally threatened speciesare notrecorded throughout (see Tana, withseventhreatened speciesinLakeTana itself e X\ XX k_\ Æ KXeX CXb\ è ©SARAHDEPPER

Jg\Z`\j`ek_\jgfkc`^_k shes listedasRegionallyExtinct(tonorthern cient species).Moreover, insometributaries CXY\fYXiYlj cientlysurveyedtoaccurately – 8le`hl\jg\Z`\jÕ Zfdgc\o ). Although et al. fZb

Victoria are stillquite evident,andit ispragmaticto record resurgence, thethreats tomany ofthespeciesinLake compared todetritivores (Witte of thespecies,withgreater resurgence ofzooplanktivores has beensupportedbyevidence ofaresurgence inseveral caution inclassifyingtheLake Victoria cichlidsasextinct extinct, duetoalackofdata. HarrisonandStiassny’s which couldnotbeclassifi and 30species(HarrisonStiassny1999:table11), as suchbecauseofinadequatesurveyingandsampling, table 10)thatmightbeextinctbutcouldnotconfi listed another54species(HarrisonandStiassny1999: the specieshadnotbeenscientifi because ofcomplicationsintheirtaxonomy(inmostcases, that mightbeextinctbutcouldnotconfi table 9)listed48speciesofcichlidsfrom LakeVictoria as ameasure ofextinction.HarrisonandStiassny(1999: which couldnotbeextrapolatedtothewholelakeandused 11-year periodforasmallpartofthelake(MwanzaGulf), because Witte suggest thatmanyofthesespecieswere actuallyextinct, cichlids; nevertheless,theybelieveditwaspremature to changes thatrepresent aseriousthreat totheendemic was undergoing catastrophic ecologicalandlimnological lake. HarrisonandStiassny(1999)recognised thatthelake disappeared orwere threatened with extinctionwithinthe 200 speciesofhaplochromine cichlidinLakeVictoria had 1). Witte et al Seehausen 1996; 1993; SeehausenandWitte1995;Oijen1996; 1992; Witte references seeHarrisonandStiassny1999;Kaufman and deforestation (forfurtherdiscussionandextensive resulting from increasing lakeside agriculture, urbanisation, fi oxygen levelsinthelake’s waters;overfi hyacinth ( predator, Nileperch ( threats, including:theintroduction ofthepiscivorous lake), resulting from acombinationofwell-documented species (81species;44%oftheassessedin Lake Victoria hasthenexthighestnumberofthreatened spawning habitatswithintherivers(Tweddle 1996). where theycongregate duringmigration,andbydegraded also threatened byfi migrate from thelakeintoaffl Thieme recover slowlyfrom populationdeclines(Ribbink2001; vulnerable tohabitatdegradationandexploitation, and produce smallnumbersofoffspring, are extremely (intra-lacustrine endemism).Thesecichlidsgrow slowly called mbuna,haveparticularlyrestricted distributions when nextre-assessed. Therock-dwelling cichlids,often some maybedowngradedtoalowerRedListcategory species forittobeassessedasthreatened, itispossible sh poisons;andhabitatdeteriorationeutrophication . 1997;Witteetal.,2007)(alsoseeChapter1,Box et al et al Eichornia crassipes Eichornia . 2005).Thepotamodromous speciesthat (1992b)initiallyestimatedthatasmany et al et al . 1992a,b;KaufmanandOchumba (1992b)were usingdatalimitedtoan sheriesoperationsattherivermouths Lates niloticus ed asprobably orpossibly ) whichhasreduced lightand uent riverstospawnare et al et al cally described).They . 2007).Evenwith this . 1997a,b;Kaufman ), andthewater shing anduseof rmed assuch rmed CHAPTER 3 | FISH 59 – sh Garra shes, Labeo currie, Barbus , a Critically Endangered , a Critically Endangered , Tilapia cessiana and T. cessiana and T. , Tilapia Jg\Z`\jèk_\jgfkc`^_k

© MELANIE L.J. STIASSNY (CR), from Kinsuka rapids near (CR), from Kinsuka rapids is Critically Endangered for the same is Critically Endangered CR), a small riverine pelagic distichodontid j_Y`f[`m\ij`kp1n\jk\ie8]i`ZXËjZiXk\i is Critically Endangered due to the impacts of due to the impacts is Critically Endangered © TIMO MORITIZ Scriptaphyosemion cauveti ), and several Endangered species, especially in the ), and several Endangered ). Many of these species are Endangered or Critically Endangered ). Many of these species are sh sh trade. Just north of the Niger Delta, in a tributary of sh trade. Just north of Neolebias powelli ( characiform, is endemic to a very localised part of the Lower Niger Delta, where it is threatened by oil exploration within the delta. trewavasae tin mining. found in the coastal species are At least 10 threatened include some drainages of Sierra Leone and Liberia; these species (e.g., Critically Endangered carcharinoides, Epiplatys ruhkopfi coffea species are vicinity of the St. Paul and Lofa rivers. These by habitat degradation caused by deforestation threatened the catfi and mining. In the Konkouré River in Guinea, Synodontis dekimpei of Guinea, the In the Fouta-Djalon ecoregion reasons. killifi is threatened a tributary to the Kolenté River, species from by expansion of the nearby city of Kindia. the from recorded species are six threatened Twenty ecoregion Equatorial Crater Lakes freshwater Western of south-west at the border and the river drainages nearby, and Nigeria (see Cameroon Teleogramma brichardi Teleogramma Kinshasa, D. R. Congo. fi Bauchi plateau, the cyprinid the Benue River on the Endangered, and the majority are cichlids endemic to the majority are and Endangered, also several killifi are crater lakes, although there :Xlc[ifej]fiÔ cXb\j , et sh cant and the , assessed as , assessed as . 1995) (especially et al Teleogramma brichardi Teleogramma 3.3.4), cient (see section . Six threatened species . Six threatened Neolebias powelli (assessed as Vulnerable) is (assessed as Vulnerable) Brycinus brevis Marcusenius brucii cation of Lake Victoria’s cichlids into Lake Victoria’s cation of . 2007). Fundulopanchax powelli et al sh . 2011). However, further collections are necessary to further collections are . 2011). However, Vulnerable); however, the small red-eyed tetra, an alestid, tetra, the small red-eyed however, Vulnerable); Arnoldichthys spilopterus extensive harvesting for the aquarium by an threatened There are 12 threatened species in Lake Tanganyika (5% species in Lake Tanganyika 12 threatened are There in the Red List of the total number of 245 species recorded species assessments). The overall number of threatened are there is lower than in Lake Malawi partly because the proportion in Lake Tanganyika; fewer species present to the total number) is lower species (relative of threatened and Malawi because the threats to Lakes Victoria compared localized (Cohen generally more are species and the Tanganyikan to Lake Victoria); compared into areas tend to have a wider distribution, extending fewer threats. are there where the African Rift lakes of of the large Outside the area with high numbers of threatened the regions Valley, the rapids in the Lower species occur in and around Congo and some coastal basins in western Africa and to 24 threatened Lower Guinea. The Lower Congo has up of Inga, and has one Critically species just upstream species of cichlid, Endangered to the Kinsuka rapids near Kinshasa. restricted apparently by the impacts threatened This species is increasingly of urbanization at Kinshasa and Brazzaville (Stiassny al distribution of this species. Several establish the precise found, especially in the species are other Endangered vicinity of Malebo Pool. found in the delta region species are At least 13 threatened including two Critically Endangered of the Niger River, by the impacts oil exploration threatened species that are in the delta, the distichodontid killifi as well as agricultural and urban development Vulnerable), (e.g., the mormyrid trophic guilds may offer a pragmatic, short-term solution a pragmatic, short-term solution guilds may offer trophic (Witte in the species rich lower Ogun River nearby, recorded are mainly threatened at Lagos lagoon. These species are (e.g., by deforestation them in categories of high threat (52 species are Critically Critically are (52 species threat of high in categories them IUCN’s noted in also are of these and many Endangered, fi Nevertheless, many as Possibly Extinct). database Defi Data lake remain species in the required and sampling are extensive surveying and more conservation to fully assess the Lake Victoria throughout Victoria Because Lake cichlid species present. status of the a signifi this represents area, lake by largest is Africa’s for a would be the requirement there challenge. Moreover, than currently number of highly trained taxonomists larger of the 600 or more to identify exist in Africa, or elsewhere, many thousands of specimens that species amongst the light of this taxonomic impediment, would be collected. In an ecological classifi 60 CHAPTER 3 | FISH system, the south,in Tradou catchment oftheBreede River water abstraction causedbyintensivefarming. Slightlyto introduced species,as wellasdeteriorationinhabitatand are threatened bycompetitionwith,andpredation by, © TIMOMORITZ Africa–vanishingislandsofsylvanfi western in remnants Benin (seeSpeciesintheSpotlight:Forest Denticeps clupeoides :``Xc<[e\\def]fJl_ 8]i`ZX and anundescribedspecies of Jflk_ ]ifd dèefn Lake Kivu, Lake Malawiare endemic, andmore than95%oftheLake in theselakes.Forexample, 99% ofcichlidspeciesin due totheendemicspecies-rich fl range speciesfoundintheRift Valley lakesare largely other partofthecontinent.Thehighnumbersrestricted many more restricted rangespeciesthanare foundinany areas; seebelow).Thelargest oftheRiftValley lakeshave (although thisislikelytobeanunderestimate insome have onlyonetothree restricted rangespecies recorded Province ofSouthAfrica)(Figure 3.7).Mostsub-catchments Africa(particularlyintheCape andsouthern in eastern Lower Guinea,andsomepartsoftheCongoBasin, several Africansub-catchments,butmainlyinUpperand distribution rangesoflessthan50,000km Restricted rangespecies(identifi 3.3.3 RestrictedRangespecies throughout muchofitsrange. widespread introduction of Highveld ecoregion. Thisspeciesisthreatened mainlybythe Zambezi-Luangwa ecoregion andpartsoftheZambezian mortimeri Oreochromis Endangered specieswithrestricted distributions,thecichlid aid drainageforagriculture. UnlikemostotherCritically to theSiandaRiverthathasbeencanalised,probably to Zambezi fl in theZambezianLowveldfreshwater ecoregion. Intheupper edgeoftheBanhineNationalParkinMozambique, eastern Banhine) isknownfrom fourneighbouringsitesatthesouth- and odli ecoregion, oodplain Barbus ), where theyfacesimilarthreats totheabove- Tilapia guinasana . Two CriticallyEndangered species andtheMalotiminnow, is widely distributed in the Middle iswidelydistributedintheMiddle O. niloticus et al Neolebias lozii ed asthosespecieswith sh, , whichoccursnaturally ocks ofcichlidsfound . (2009)givesashort Barbus , which is displacing it , whichisdisplacingit Clarias cavernicola ( Pseudobarbus ² Pseudobarbus Barbus ) are foundin is restricted isrestricted sp.nov. , , CHAPTER 3 | FISH 61 © JOHN FRIEL Lake Tanganyika Lake Tanganyika contains many restricted range cichlid species, 95% of which are endemic to the lake. Africa. continental mainland across sh species – CXY\fYXiYlj Jg\Z`\jèk_\jgfkc`^_k

ock of endemic cyprinids of (see fZbèCXb\KXeXÆk_\ sh fauna of Lake Turkana is composed sh fauna of Lake Turkana Labeobarbus ). Zfdgc\o species in the crater lakes are also mainly cichlids. Many species in the crater lakes are range species, which also of those lakes with restricted species, are and Endangered hold Critically Endangered potential candidates for designation as Key Biodiversity Extinction sites (see Alliance for Zero or possibly Areas, chapter 8, section 8.3). has nine to the north-east of Lake Victoria, Lake Turkana, lakes of Unlike the other large range species. restricted the fi the Rift Valley, mainly of nilotic riverine species rather than cichlids. Lake range species, mostly has eight restricted in Ethiopia Tana by the species fl represented the genus Figure 3.7. The distribution of restricted range freshwater fi range freshwater of restricted distribution 3.7. The Figure sub-catchment. per river/lake of species = number richness Species 8le`hl\jg\Z`\jÕ 62 CHAPTER 3 | FISH practising catch and release. practising catchandrelease. is promotedasa fl the middleandlower Vaal and Orange rivers,respectively. It common,especiallyinlargedeeperpools it isreasonably is endemictotheOrangeRiversystem inSouthAfrica,where The largemouthyellowfi cichlid speciesendemictoLakeTanganyika.A selectionoftherockyshore they are adaptedtothefastcurrents, lowlightintensity, Endangered, mostlikelyshowrestricted rangesbecause ecoregion. Thesespecies,someofwhichare assessedas of Inga,inthelowerpartLowerCongoRapids Up to17restricted rangespeciesare foundinthevicinity range species,thegreatest numberbeingintheCess. John, andCessrivershavebetweenfi the relatively smallcoastalbasinsoftheLofa,St.Paul, restricted rangespecies.IntheUpperGuineaprovince, A numberofriversalsohavenotablyhighernumbers agship anglingspecies, withmostanglers sh, Labeobarbus kimberleyensis © SAIAB/ROGER BILLS veand12restricted (NT), range species are: theupper partoftheTana River, near sub-catchments withthegreatest numbers ofrestricted generally lower numbersofspeciesfound there. The restricted rangespecies, possiblyaconsequenceofthe Africahave fewer andsouthern The riversofeastern of theOgowe,andinlowerSanaga River. (fi province, thegreatest numbersofrestricted rangespecies (all withfi in theUpperLualabaandBangweulu-Mweruecoregions and inseveralpartsoftheLufi species), theupperpartofLowaBasinnearLakeKivu, (six species),theLubibasindrainingtoSankuru(seven range speciesincludetheUbangiRiverinregion ofBangui parts oftheCongoBasinwithhighnumbersrestricted reason asfortheLowerCongorapids(seeabove).Other compared tosurrounding areas, probably forthesame higher numbersofrestricted rangespecies(sevenspecies) The UpperCongorapidsecoregion alsohasmarginally Lower Congorapidsregion assurveysthere continue. that more restricted rangespecieswillbefoundinthe (Lowenstein identifi in theLowerCongoisincomplete,butrecent research has knowledge ofthetaxonomyandbiogeographyspecies complex atInga(seesection3.3.2).Thecurrent stateof be impactedbydevelopmentofthehydropower dam and highturbidityoftherapids.Theyare alsolikelyto ve to eightspecies)are foundintheIvindoandupperparts ed manymore species than were previously known ©SASKIAMARIJNISSEN ve restricted rangespecies).IntheLowerGuinea et al . 2011;Stiassny ra, Luvira,andLuapulabasins et al . 2011).Itisprobable CHAPTER 3 | FISH 63 ow into cient (DD) (Figure 3.8) are 3.8) are cient (DD) (Figure ve restricted range species, range species, ve restricted cient information to make a reliable cient information to make a reliable cient species cient ve restricted range species); the Ruvu River near Ruvu River the range species); ve restricted Lake Tanganyika (seven restricted range species). The range restricted (seven Lake Tanganyika in southernonly catchment with moderately high Africa River in species is the Olifant range restricted numbers of that holds fi Cape the Western total number of species 75% of the around representing catchment. in the recorded 3.3.4 Data Defi Data Defi Species assessed as uncertain, or for taxonomy remains those for which the is insuffi which there Nairobi (fi Nairobi the and range species); (eight restricted Salaam Dar es near its outfl of the Malagarasi River lower part mainland continental Africa. Species sh species across fi cient freshwater © ROBERT SCHELLY © ROBERT Figure 3.8. The distribution of Data Defi Figure richness = number of species per river/lake sub-catchment. richness = number of species per river/lake The Lower Congo rapids, home to many restrictedThe Lower Congo rapids, specialist species. 64 CHAPTER 3 | FISH town of Bangui(11Data Defi Defi species), theUpper Congorapidsecoregion (15Data downstream intheLower Congo(13to19DataDefi around MaleboPool andsub-catchmentsimmediately parts oftheCongoBasin,withhigher numbersparticularly Five ormore DataDefi taxonomic problems abound. the endemiclakecichlidsisnotoriouslydiffi lacking. Anotherimportantfactoristhatthetaxonomyof distribution andecologyofmanythesespeciesisstill have notbeenwellsurveyed,anddataonthecomplete species numbers(seesection3.3.1),extensiveareas a combinationoffactors.Thelakesare richinoverall Data Defi being classifi of theselakes,witharound 35%oftheassessedspecies the highestproportion ofDataDefi in theRiftValley Africa.LakeVictoria lakes ofeastern has The greatest numbersofDataDefi African freshwater fi defi of DataDefi this underrepresentation, themapshowingdistribution map thedistributionofthesespecies.Evenallowingfor with certainty. Insuchcasesitmaybeimpossibletoreliably a speciesinparticularriversystemcannotbedetermined locality information,suchthatthepresence orabsenceof collections isthattheyfrequently containambiguous A persistentproblem presented byolder, historical revealed torepresent speciescomplexes. scrutiny andwithbroad geographicsampling,willlikelybe putatively ‘widespread’ speciesthat,onclosertaxonomic 2004). Thisistruefornumerous otherpoorlydiagnosed, have yettobediagnosedanddescribed(Tweddle but itprobably represents acomplexofspeciesthat widespread acrossandcentralAfrica, muchofsouthern good caseinpoint.Ascurrently defi as anythingbutDataDefi complexity precludes assigningtheconservation status many speciesof specimens thattheirtaxonomyisunresolved. There are Some speciesmaybesopoorlyrepresented bycollected type localityfrom where thespecieswasfi 25% to80%were knownonlyfrom thetypeseriesor found that,forthedifferent generaandfamiliesselected, of Africa(CLOFFA, Daget mormyrids includedintheCheckListofFreshwater Fishes Stiassny (1999)tookarandomselectionofcatfi from onlyoneorafewspecimensfrom asinglecollection. the totaldistributionofspecies.Manyspeciesare known reason forthisistheabsenceofreliable informationabout freshwaters intheirsub-catchment.Themostcommon assessment ofhowtheyare impactedbythreats tothe ciency isasignifi cient species),andtheUbangiRiver adjacenttothe cient species foundintheselakesresults from cet fi cient ed asDataDefi Barbus sh speciesclearly indicatesthatdata shes. cant problem forassessments of cient species are foundinseveral , forexample,where taxonomic et al cient. cient. Thehighnumbersof cient species). Thehigh . 1984,1986,1991),and cient speciesare found Barbus eutaenia cient speciesforany ned, cult, andmany rst described. rst B. eutaenia shes and shes isa cient et al is . Data Defi Tana systeminKenya coversalarger area and hasmore species), andin theGalanabasin(inKenya).However, the Basin attheborder between Tanzania andKenya(seven of DataDefi In theEastCoastprovince there isasmallconcentration explored andpoorlyknown areas inAfrica. province describedbyRoberts(1975))isoneoftheleast (which roughly corresponds withtheQuanza ichthyofaunal 10 DataDefi tributary tothemiddlesectionofQuanzathathas catchments withfi are usuallylow–theexceptionsbeingacoupleofsub- Lower Guinea,thenumbersofDataDefi freshwater ecoregions.Africaand But,aswithwestern catchments oftheQuanzaandZambezianHeadwaters Data Defi Defi of theRokelBasininSierraLeoneeachhavefi middle partoftheKonkouréBasininGuineaandmost Africa,the Republic ofCongo(Brazzaville).Inwestern Niari-Kouilou system(fi Data Defi in Cameroon, thelowerreaches oftheIvindoRiver(fi species) andtheKribiRiver(fi the middlereaches oftheNyongRiver(sixDataDefi province thatare noteworthyforDataDefi centralAfrica.OtherpartsoftheLowerGuinea of western River wasthoughttobeoneofthebetter-surveyed rivers was aparticularsurprisebecause,priortothis,theCross previously beenunderestimated byasmuch73%.This found thatthenumberofspeciesinCross Riverhad many ofthesespecies.Forexample,Teugels morea needtolearn aboutthedistributionandecologyof ichthyofaunal provinces (Reid1989),andthere isevidently interesting, sinceitincludesamixofspeciesfrom thetwo Data Defi the headwatersofCross inCameroon containnine Nilo-Sudan andLowerGuineaichthyolofaunalprovinces, drainages freshwater ecoregion, attheborder ofthe few exceptions,however. GulfofGuinea IntheNorthern Africa,butrarelyof western inlarge numbers.There are a centralAfrica,andinseveralsub-catchments of western through mostoftheLowerGuineaichthyofaunalprovince Data Defi contain more thanfi parts oftheBlueandWhiteNilesysteminSudan extent, fortheNilesystem,where LakeTana andsome and ecologymaybelacking.Thisisalsotrue,toalesser collected from theseregions, butdataontheirfullranges Africa, relatively large numbersofspecieshavebeen well explored. AswiththeRiftValley lakesofeastern Congo Basintendtobeinareas thathavebeenrelatively numbers ofDataDefi cient species. cient cient species(fi cient species) inGabon,andalarge partofthe cient species.Thisregion isbiogeographically cient speciesare alsofoundinseveralsub- cient speciesare foundinsub-catchments cient speciesintheheadwaters ofthePangani cient species.Theentire Quanzaecoregion ve DataDefi ve DataDefi cient speciesrecorded from the ve DataDefi ve tonine,withup to32%ofthe ve DataDefi cient species. cient cient species,andone cient species)inthe cient speciesare cet species) cient cet species cient et al v Data ve . (1992) cient ve CHAPTER 3 | FISH 65

); e c

was cally cient. . (2003), et al Salmo pallaryi Aplocheilichthys ; c descriptions. This is Barbus microbarbis sp. nov. ‘Naivasha’ is a sp. nov. © JEAN-PIERRE BOUDOT ed as Extinct ( . 2011). Without a full scientifi Barbus microbarbis et al Aplocheilichthys but, according to Seegers but, according A. antinorii known from its type locality, Lake Luhondo (=Ruhondo) locality, its type known from in Rwanda, though it possibly also inhabited the small all three had restricted distributions and their apparent distributions and their apparent had restricted all three at least in part, to introductions attributed, extinctions are of alien species. poeciliid of indeterminate taxonomy that has been reported as been extinct since the is quite distinct. It has probably from 1970s or 1980s, following competition or predation The cyprinid species. introduced Some species have not been formally, scientifi Some species have not been formally, by taxonomists nonetheless recognized described, but are not formally described they are as being valid. The reason had the time or is usually because taxonomists have not to publish the scientifi resources and capacity support for more need evidence of an urgent taxonomists in Africa (Stiassny building for freshwater 2002; Lowenstein of the species description and account of the distribution status, it is not possible to ascertain their conservation categorized as Data Defi and they must remain and some of exist in the literature, examples Numerous but, here in the analyses presented included these are not usually Red List guidelines, they are to the according thought likely added to the IUCN Red List unless they are to be threatened. 3.3.5 Extinct species classifi species are Three ‘Naivasha’; sp. nov. sh sh species. cient species cient cient species; cient in some parts of cient in some nd large numbers of fi nd large . 2011), and some of these catchments are . 2011), and some of these catchments are et al cient species, but these represent the majority of cient species, but these represent ooding. Such conditions present challenge to the freshwater a considerable these habitats. The region species that live in is in undisturbed parts of the Congo forest where one would where in undisturbed parts of the Congo forest to fi expect reasonably A similar situation exists for the Sudd wetland in southern Sudan. Lack of data in these cases is undoubtedly due a need indicating an urgent lack of surveys in these regions, or undescribed rare surveys before to undertake targeted (see Chapter 8). extirpated species are species. A lack of species is unsurprising in the Sahara, such as the Kalahari and and several other arid areas in southern Namib ecoregions Africa, and the Shebelle- eastern in Juba ecoregion Africa (as discussed in section also sub-catchments in parts are there 3.3.1). However, of the Congo main basin that have no assessed species (Stiassny assessed species being Data Defi rivers of the the basin). This illustrates that, although most low characterized by relatively are East Coast province is required research species numbers, considerably more for several of these species. and The catchments in the Atlantic Northwest have ecoregions Mediterranean Northwest freshwater consistently low numbers of Data Defi Defi in all these catchments, the Data however, (and in some of the species present 25% or more represent cases up to 100%). Similar patterns seen in the Horn are of Africa, and in the south-western Cape of Africa, where two sub-catchments with only one or numerous are there Data Defi also are the assessed species in those catchments. There fi is a complete lack of data for any there where areas home to an endemic group of ’Maghreb barbs‘, which are increasingly threatened habitat. by loss of Rivers in northern Africa typically have a hydrological regimeunpredictable that is and which may experience periods of intens fl 66 CHAPTER 3 | FISH dioxide contained under pressure indeepwaters and occasional fi commercial developmentoftheregion fortourism,and water abstractionfortheneighbouring townofKumba, agricultureand-burn andoil plantations, deforestation, These includesedimentation and pollutionfrom slash- crater lake,Barombi Mbo,faceseveralseriousthreats. (GCCA Forum2007).Nevertheless, thefi was exportedin2007totheUSAforaquariumpurposes (Reid 1991).However, thespeciesisstillextantbecause it be extinct,becausefi mongo Harrison andStiassny(1999)thoughtthecichlid be affected byaquaticweeds. inadequately treated humanwaste.Thespeciesmayalso to waterpollutionfrom agriculture, andpossibly alsofrom the impactsfrom thedams,habitatqualityisdecliningdue to resolve whetherthespecieshasdisappeared. Besides it (DeVos 1995).More concertedsurveyingisnecessary had anameforit,butcouldnotconfi least priorto1995,localfi Akosombo damsontheVolta Riverinthemid1960s.At to modifi and itispossiblethatthespeciesbecameextinctdue made tocollectthisspeciesbetween1961and1988, note thatintensivebutunsuccessfulattemptswere collection before 1943.HarrisonandStiassny(1999) been confi Endangered intheRedList.However, there havenot the lowerVolta Basin andiscurrently categorizedas The schilbeidcatfi the recent IUCNRedListassessments. other speciesthatare notclassifi discussed thepossibleevidenceofextinctionforseveral Harrison andStiassny(1999)Helfman(2007)have of theintroduction ofspecies valid speciesitisalmostsurely extinct,possiblyasa result (De Vos caught in1934,despiteintensivesamplingtheregion streams fl Sidi Ali,mightbeconspecifi verte’) from LakeIsli,tothesouth-westofAguelman de (2005) notethatanunnamedpopulationoftrout (‘truite of commoncarpin1934,althoughDellingandDoadrio extinct around 1938,probably duetotheintroduction (Delling andDoadrio2005).Thespeciesapparently went Morocco,northern andknownfrom atleast19specimens de SidiAli,ahighaltitudelakeintheAtlasMountainsof The salmonid, between bothgenerahavebeenidentifi be correct because,inotherregions, putativehybrids Barbus suspected tobeahybridbetween (De Vos , endemictoLakeBarombi Mbo, species(Banister1973).Thishypothesiscould et al cation oftheriverfl et al owing intothelake.Onlyonespecimenwasever rmed reports ofthisspeciessincetheoriginal sh killscausedby sudden releases ofcarbon . 1990).However, thesoletypespecimenwas . 1990;HarrisonandStiassny1999).Ifitisa Salmo pallaryi sh Irvineia voltae Irvineia shermen hadnoteditsabsence shermen knewthespeciesand c with c ow afterconstructionofthe , wasrestricted toAguelman Tilapia ed assuchaccording to S. pallaryi rm thattheyhadseen isknownonlyfrom ed (Wamuini 2010). Varicorhinus and

Cameroon, might . Haplochromis shes ofthe Stomatepia anda

soils, which are easilyeroded, are exposed andwashed in signifi Africa(Stiassny western of riversystems(andlakes,such asthecraterlakesof deforestation cansignifi 2009; Brummett these DDspecies. continent mightreveal evidenceofextinctionforseveral expect thatfurtherresearch andsamplingthroughout the the continent(seesection3.3.4),itisreasonable to high numbersofDataDefi species recorded from theseregions: seesection3.3.2)and Africa (asindicatedbythelarge numbersofthreatened Given theabundantthreats tospeciesinmanypartsof scope ofthisundisputeddeclineincichlidspecies. in order tofullyunderstandthegeographicandtaxonomic for more research andsurveyingofLakeVictoria fi categorized asDataDefi noted insection3.3.2,manyofthespecieslakeare or theirtaxonomyisnotsuffi Extinct becausethere hasbeeninsuffi but manyofthesecannotbedefi cichlids endemictoLakeVictoria thatare possiblyextinct, lake sincethe1980s.There are indeedmanyspeciesof Victoria, withthedeclineofendemiccichlidfi historical times(since1500AD)mayhavebeeninLake It isoftennotedthatthelargest extinctioneventinrecent lake, iscurrently considered tobeCriticallyEndangered. (Stiassny in 1986,killingmore than5,000peopleandlivestock) sediments (similartotheeventthatoccurred atLake Nyos communities withinthecatchments (Bradshaw impacts thefreshwater speciespresent andthehuman riparian forest canaffect peakfl allochthonous materialsfrom theforest. The removal of and theCongoBasin,results infoodwebsdependenton the riversinforested partsofUpperandLowerGuinea, dissolved mineralandnutrientconcentrationofmany then promote algalbloomsandeutrophication. Thelow al temperature andhydrological regime ofrivers(Brummett fi 2010). Lossofforest coverdeprivesmanyspeciesof between tree-covered andopenareas (Ibala-Zamba shoreline, speciescompositionwasfoundtodiffer clearly the Léfi even onamicro scale,hasbeendemonstratedrecently in threats tofreshwater fi increased sedimentation,isoneofthemostwidespread Habitat modifi Deforestation, habitatlossandsedimentation 3.4 Majorthreats tospecies shes ofshelterfrom predators, andchangesthewater . 2009).Excessivesunlightandhighertemperatures may ni River, where withinarelatively smallstretch of cant increases insedimentation asdelicateforest et al cation, through deforestation andassociated . 2011). et al Stomatepia mongo . 2009;Farrell shes inAfrica.Theeffect ofthis, cient,andthere isanurgent need et al cantly affect theecohydrology cient speciesfoundthroughout . 2011)).Deforestation results ciently welldescribed.As ow fl ow nitively categorizedas ooding events,which et al , endemictothis cet sampling cient . 2010).Thus, shes inthe et al sh fauna, . 2007, et CHAPTER 3 | FISH 67 et shes . 2011). In . 2005), with et al . 2011). Mining et al ve months in most et al shes in the Malagarasi cantly contributed to habitat cantly . 2005). Similarly, the high number the . 2005). Similarly, 2009). . 2009; Brooks . 2009; Brooks et al et al. et al . 2011) and Upper Guinea, war and civil cation. et al © K.-D.B. DIJKSTRA . 2009; Smith unrest have displaced local communities into previously local communities into previously have displaced unrest with these communities of forest, undisturbed regions often settling along waterways (Thieme loss and high levels of sedimentation in many regionsloss and high levels of across Africa. al and Kasai is especially common in the Congo, Sanga, impacts riparian basins. Land conversion for agriculture as well as many other habitats. In eastern Africa, forests, fi the high number of threatened into the of agriculture Basin is due to encroachment wetlands (Darwall draining to the species in the Ruvu River, of threatened many regionally from results probably coast of Tanzania, at high risk of endemic species being found in areas habitat modifi land, causing placed on the cleared frequently Livestock are of land poorly managed processes and these overgrazing, and in accelerated soil erosion clearing and grazing result The Ubangi River in central Africa sedimentation. increased mining and by sedimentation from impacted is so severely river of Mpoko that the reduced region in the agriculture for four or fi shipping depth prevents years (Brummett loss of riparian habitat and deterioration Not surprisingly, of high in areas greatest are ecosystems of freshwater human settlement. For example, several species of fi loss in the heavily populated lower impacted by habitat are Ogun Basin in Nigeria, as well as in the northern parts of cities of Kinshasa Lower Guinea, and close to the large and Kisangani in the Congo Basin (Stiassny several parts of the Congo Basin (particularly the eastern part of the basin and in the vicinity of the Sangha; see Brummett cover. consequent loss of forest Mining, such as observed here in the East Nimba Forest Reserve has signifi in Liberia, many regions. loss and high levels of sedimentation in mines, and commercial Small-scale alluvial mining, larger rivers basins in extraction of sand or clay have impacted parts of southern, western, and central Africa (Darwall et . 2009; . 2009). et al et al cally for the 2011). Deforestation is a 2011). Deforestation shes and suffocate their shes and suffocate et al. . 1998). The Kasai, Sanga, and sh populations. The increased The increased sh populations. cation of logging and agriculture of logging and agriculture cation et al cantly contributed to further habitat cantly rewood and charcoal is a more serious serious is a more and charcoal rewood . 2011). Deforestation has also taken place . 2011). Deforestation . 2011). et al et al . (2009) note that in western. (2009) note that in the Africa (which includes shes in the Upper Guinea region that are impacted by impacted that are Guinea region shes in the Upper specifi Deforestation and charcoal. rewood Deforestation for logging also opens up large tracts of logging also opens up large for Deforestation for example, by mining and for further exploitation, forest bush meat hunting, and settlement. ventures, agriculture Mining has signifi al ichthyofaunal provinces), Upper Guinea and Nilo-Sudan of along the banks is especially prevalent deforestation and Senegal rivers. Some examples of Niger, the Volta, fi discussed in section 3.3.2. The central are deforestation lost an estimated 46% of its has already African region and to logging and conversion to agriculture, rainforest watershed at an average rate of continues to lose forested 7% per year (Revenga in many parts of the Lower Guinea province, major threat (e.g., in the and much of this is associated with logging the Ogowe/Ivindo system (Gabon), Nyong (Cameroon), However, and Kouilou/Niari systems (Republic of Congo)). production, as well as slash-and-burn charcoal agriculture, other major are and oil palm plantations banana, rubber, and associated sedimentation drivers of deforestation and in Lower Guinea (especially in Cameroon), through some parts of the Congo Basin (Brummett along the Cameroon rainforest biodiverse Extremely the Ndian to the Kribi (Kienké) rivers, from coast, roughly has been converted to oil palm, as have parts of the Upper plans for are and there ecoregion, Congo freshwater Lake Tumba development of oil palm plantations around (Brummett to allow for the development of eucalyptus plantations along the coast of the Republic of Congo and Cabinda. Eucalyptus is mainly used for building materials and for fi of fi production of western for parts problem and central Africa, especially riparian remnant development, where of urban near areas (Smith vegetation is particularly under threat Brummett turbidity can clog the gills of fi clog the gills of turbidity can levels the light reducing 1993), as well as eggs (Roberts so photosynthesize and plants cannot so that submerged of the waters. eutrophication die, exacerbating Expansion and intensifi from Threats common causes of deforestation. are in and widespread particularly strong are deforestation and in the Congo Guinea provinces, the Upper and Lower encompass the last remaining Basin, given that these on the continent. Smith regions extensively forested some of the more are ecoregions Upper Congo freshwater in terms of ongoing deforestation, seriously impacted areas deforested while the Lower Congo has been almost entirely for many decades (Stiassny into streams, rivers, and lakes. The sediment covers covers The sediment and lakes. rivers, streams, into for breeding habitat suitable reducing surfaces, submerged of many fi and feeding 68 CHAPTER 3 | FISH et al studies (Darwall freshwater systemshavebeenreported inallregional impacts ofpesticidesandfertilizersfrom agriculture on Water pollutionisaproblem inmanypartsofAfrica.The Pollution section 1.2.2.1). by 2025(seeBrummett development on thelowersectionofCongo River such asthe39,000MW‘Grand Inga’damproposed for is alsolarge, includingplans forsomeverylarge dams, for future development ofhydropower projects inAfrica for theAswanDam(seesection 3.3.2).Thepotential The overallimpactofdams may besevere, asnoted also unsuitableforhabitationbypreviously nativefi extent thatthehabitatsimmediatelybelowdamare fl Africa. Downstreamenvironment, especiallyinsouthern lacustrine speciesthatmaybecomeinvasiveinthelake the attentionofanglersforintroduction ofexotic originally present. Theselakereservoirs alsoattract uninhabitable formanyoftheriverinefi of fi Africa. Damspreventsouthern thelongitudinalmigration Africa,andZimbabwein Africa,muchofwestern northern and someofthelargest, are intheMaghreb province of 1, Figure 1.3).Thegreatest concentrationsofdams, large dams(>500,000m large riversofAfrica;atleast135theseare classifi Some 1,207damshavebeenconstructedonsmalland Dams the RepublicofCongoandAngola. species incoastalfreshwater systemsofGabon,Cabinda, species intheNigerDelta,andmayposeathreat to habitat, specifi Pollution from oilexploration,andassociatedlossof those closetolarge cities,suchasKinshasaandLagos. traffi other urbanindustries,carsinthecities,andfrom boat settlements. Pollutionfrom oilexploration,factoriesor threatens fi Organic pollutionfrom humananddomesticwaste problem incentralAfrica. of suchpesticidesasamethodfi compromising theirphysiologyandbehaviour. Theuse al Africa(Smith trypanosomiasis (e.g.,inpartsofwestern programs fordiseaseslikemalaria,schistosomiasis, and prevalent (seeabove).Theuseofpesticidesinvectorcontrol the regions where small-scaleorcommercial miningis (see above)).Pollutionfrom miningisaseriousthreat in Malagarasi Basin,whichisimpactedbyagriculture result ineutrophication ofthelakesandrivers(e.g., sedimentation causedbysoilerosion, andfrequently are usuallyalsocoupledwithimpactsfrom increased ow andsediment loadmaybechangedtosuchan . 2009))mayalsoimpactfi sh, andcreate lake-likeconditionsupstream thatare c onriversimpactsthefreshwater systems,especially . 2010a;Brooks shes inallareas where there are sizeable cally threatens severalrestricted range et al . 2005,2009;Smith et al. ³ et al ) (FAO 2010)(seealsoChapter 2011).Theeffects ofpollution shes, withlowdosespossibly . 2011)(butsee Chapter1, shing isanadditional et al shes thatwere . 2009;García ed as shes. et © WILLIAMDARWALL to anyendemicspecieswhoserangeisrestricted tothe fl water removal signifi to around 1,350km surface area ofthelakefrom 25,000km impacted thewetlandsofLakeChad,greatly reducing the abstraction (coupledwithclimatechange),hasseverely Africa,water Africa (seesection3.3.2).Inwestern andsouthern have humansettlements,suchasnorthern Water abstractionisathreat particularlyinaridareas that homogenization ofthepopulations. normally isolated.Thismayresult indetrimentalgenetic possibility ofinterbreeding betweenpopulationsthatare species tonewbasinsbeyondtheirnormalrange,withthe et al have facilitatedthespread ofintroduced species(Darwall In SouthAfrica,inter-basin transferschemesofwater agriculture, andindustrialdomesticconsumption. transfer ofwater)andabstractionwatertosupply by channelization(oftenforirrigationorinter-basin The geomorphologyandfl River channelizationandwaterabstraction approved. fornewdamsare and operationprocedures, construction, These impactsneedtobeevaluatedbefore Ghana, willhaveasignifi Dams, suchastheAkosomboDamonVolta Riverin Burkina Faso. in ontheKoustream channelization, as isthecasehere The geomorphology andfl ow ofastream orriver, thismayposeasignifi . 2009),andmayalsoallowthemigrationofnative © TIMOMORITZ ² in2001(García cantly disturbs theenvironmental species. cant impactonfreshwater ow ofmanyriversisaffectedby ow ofmanyriversisaffected et al ² intheearly1960s . 2010a).When cant threat cant

CHAPTER 3 | FISH 69

et Lates shing cation sheries . (2009) . (2009) sheries for sheries sh species et al shes in Lower sh size is accepted in sh size is accepted Arnoldichthys spilopterus . (2010b) note that are unsustainably harvested are et al shing down the food web. Allan shing down the 2011) and et al. ) to Lake Victoria in East Africa has been ) to Lake Victoria Alestes dentex and shed, particularly the Volta. Those authors report report Those authors the Volta. shed, particularly been reported sh poisons and explosives) have also . (2005) note that this decline in fi that this decline in . (2005) note sh in the cuisine. García sh in the cuisine. García Lates niloticus also note that many freshwater bodies in westernbodies in Africa freshwater that many also note overfi are species of larger has been a disappearance that there in some western a as the Oueme, as African rivers such sequentially fi of result al for small preference some regional parts of Africa, due to fi the lake into river mouths to spawn. Smith to spawn. mouths into river the lake Barbus bynnii, Hydrocynus niloticus, Anguilla anguilla, forskahlii, (overharvesting or the use of small mesh, unselective fi (overharvesting or the fi gear, such as Malebo particularly in areas for the Congo Basin, and Mai N’dombe. Some fi Pool, Lake Tumba, and artisanal fi also the focus of commercial are the aquarium trade, for example, some killifi Guinea (Stiassny (see section 3.3.2). Invasive species in many parts of Africa, a problem Invasive species are modifi and their success has been aided by habitat The impact and the development of dams (see above). Nile perch cichlids and the introduced on haplochromine ( in northern Africa. Threats from poorly managed fi poorly from in northern Africa. Threats and 3.3.5). In extensively documented (see section 3.3.2 haplochromine eastern tilapiine and Africa, introduced focused sheries are (VU), the Niger tetra, is endemic to Nigeria, where it is restricted to the lower Ogun and lower Niger sh communities, such as seen heresh communities, on © JOHN FRIEL shing © TIMO MORITIZ shing is a major threat to many species in the Great to many species in the Great shing is a major threat rivers. Arnoldichthys spilopterus The high diversity of fi are often impacted when dams are the Malagarasi, Tanzania, constructed. area where the water removal is occurring. Small dams the water removal where area abstraction may or weirs, river channelization, and water the weirs are all operate together in some places, where deeper pools from installed to divert water and create which it is easier to extract the water by pump. Overfi Overfi such as in Lake Malawi Lakes of the African Rift Valley, fi (see section 3.3.2), particularly where of species as they move from along the migration routes 70 CHAPTER 3 | FISH Pedjari National Park, Benin, pictured here. here. Pedjari NationalPark,Benin,pictured Baliwaterholein out,suchastheMare many lakesdrying Drought isaseriousproblemformanypartsofAfrica,with dolomieu European orNorthAmericanspecies(e.g., fi Invasive speciesposethegreatest recorded threat to al of Rwanda),through competitionandpredation (DeVos ruandae cichlids are themselvesathreat toacyprinid, genus X]i`\e[Xe[]f\ all ofAfrica(ChenjeandMohamed-Katerere 2006)(see and maybeasmuchUSD100millionannuallyacross at USD20-50millioneveryyearinsevenAfricancountries, The economicimpactsofthewaterhyacinthare estimated to freshwater ecosystemsinmostofsub-SaharanAfrica. invasive speciesinAfrica,presenting aconsiderablethreat ( Africa(seesection3.3.2).Thewaterhyacinth northern have hadverysignifi the forested regions ofAfrica,although theprecise nature 1, Figure 1.5).Climatechangeisalso expectedtoimpact (PietersenandBeekman 2006)(seealsoChapter patterns 50% ofthisreduction wascausedbychangesinclimate the 1960sand2001(seeabove), anditisestimatedthat Africa, wasreduced to 5.4%ofitssurfacearea between seasonal orhavedriedcompletely. LakeChad,inwestern where manyonce permanent streams havebecome Africa, serious problemandsouthern inpartsofnorthern thirds ofthefreshwater fi the secondmostseriouscauseofdeclineforalmosttwo According toGarcía natural disasters,suchasdrought, andtoclimatechange. noted aboveare especiallysusceptibletotheimpactsof Fish faunasalready weakenedbymanyofthethreats Climate changeandextreme events kfk_\]i\j_nXk\iY`f[`m\ij`kp Africa,mainlycomingfromshes insouthern introduced Eichhornia crassipes Eichhornia . 1990).

Jg\Z`\j`ek_\Jgfkc`^_k Jg\Z`\j`ek_\jgfkc`^_k Oreochromis ,

in Lake Luhondo (a small lake in the northern part in LakeLuhondo(asmalllakethenorthern ; Oncorhynchus mykiss ), andfrom invasivespeciesofthecichlid . Introduced mosquitofi cant impactsonnativespecies in et al ) isoneofthemostwidespread sh in northern Africa.Droughtsh innorthern isa . (2010b),naturaldisastersare – – K`cXg`Xè\Xjk\ie8]i`ZXÆ NXk\i_pXZèk_#Xk_i\Xk and ). Salmo trutta © TIMOMORITZ sh ( sh Varicorhinus Micropterus Gambusia ; alsosee et ) (García water abstraction,andgravelextractionfrom riverbeds parasitic pathologies,pollution,constructionofdamsand overfi this declinemaybeattributedtothecombinedeffects of Endangered) throughoutAfrica, itsrange.Innorthern decline oftheEuropean eel( A more recent exampleisprovided bythecalamitous Lake Victoria inthe1980s(seealsoChapter1,Box 1.1). resulted inthedeclineofmanycichlidspecies of Stiassny (1999)discusshowseveralcombinedthreats rather thantheresult ofanysinglethreat. Harrisonand species isaproduct ofacombinationthesefactors, cases, declineinpopulationsizeanddistributionofany impacting thefreshwater fi In conclusion,there are manytypesofthreat thatare given inChapter8,thisvolume. detailed accountoftheimpactsclimatechangeare substantially different from theexistingconditions.Amore species mayexperiencehydrologic conditionsthat are by the2050s,more than80%ofAfrica’s freshwater fi et al of theseimpactsisunclear(Schiermeier2008;Thieme reduce thetotal numberofspecies thatare classifi A majorchallenge istoaccumulatedatathat canhelp the species,and tomakesuitableconservation decisions. very diffi particular species.Intheabsence ofsuchinformationitis very littleadditionalinformation abouttheecologyofeach distributions withinacatchment, there is,however, often Where informationis availableonspecies’presence and the westintomiddleCongo(Ibala-Zamba2010). Léfi resulting from therecent studyontheichthyofaunaof Basin), iswellexemplifi fi ecological data,andtheneedforfurtherresearch onthe of knowledgeeventhemostbasicdistributionaland regions tosupportconservationmanagement.Thelack basic baselinedataforseveralpotentiallybiodiverse Considerable additionalresearch isrequired toprovide signifi also confi (see section3.6)andpriorities.However, thestudyhas and hasidentifi and conservationstatusoffi This studyhighlightsspecifi 3.5 Research actionsrequired invasive species(seesection3.3.2). groundwater abstraction,pollution,andintroduction of Africa isattributabletoacombinationoffactorsincluding saourensis shes oflarge partsofAfrica(inparticular, theCongo ni River(as reported above),atributarydrainingfrom . 2010;Brummett shing ofsilvereelsincoastalwaters,theimpacts of cant gapsinourknowledgeofthisfaunastillexist. cult toaccuratelyassess theconservationstatusof et al rmed theopinionofLundberg toCriticallyEndangered statusinnorthern . 2010b).Similarly, thedeclineof ed someconservationrecommendations ed bytheincreased knowledge et al shes throughout thecontinent, sh faunaofAfrica.Inmost ofthedistribution c patterns . 2011).Itisexpectedthat, Anguilla anguilla et al . (2000)that ) (Critically Aphanias ed as ed sh CHAPTER 3 | FISH 71 nances and nance can be nance can be rm that the freshwater cult for policy makers to set cantly threatened in many parts cantly threatened sh market on the banks of the Congo ndings of this assessment confi shing down the food web. Standardized monitoring monitoring Standardized web. the food shing down shes of Africa are signifi shes of Africa are as well as changes in biomass and local incidences of of local incidences and in biomass as changes as well fi any step before a routine be implemented as should also on or along the waterway. is developed infrastructure large political and socio-economic however, In many regions, and lack of fi logistical problems, instability, most to hamper even the expertise combine taxonomic If these monitoring programmes. basic studies, including training and fi of infrastructure, problems of studies then hopefully the recommended remediated, diversity and ecology to monitor species and population can be initiated. the health of the fauna recommendations 3.6 Conservation The fi fi to assume that even of the continent, and it is reasonable will be exerted on this fauna in the future. stress greater Nevertheless, it may be diffi as priorities when there conservation recommendations with ensuring that issues associated many other urgent are of living. The guaranteed an acceptable standard people are to meet basic human requirements many actions required of freshwater may appear to be at odds with the objectives to a need is, however, biodiversity conservation. There ll shing, to fi eld work cult for aquatic © R. SCHELLY sheries to local economies can been seen by this thriving fi culties lie mainly in the complexities of cient (514 species, 18% of all assessed species) species) assessed 18% of all species, cient (514 ows required to support this biodiversity, and additional to support this biodiversity, ows required River, at Mbandaka, D. R. Congo. River, The importance of inland fi Data Defi Data fi of prioritising discussion 8 for (see Chapter these knowledge gaps). these knowledge ecosystems on freshwater of climate change The impact an important concern,of Africa is be but one which may resilience of the a better understanding mitigated through management proper and through of species to change, 8, section 8.6.1). (see chapter resources of freshwater diversity on the further research requires this also However, the environmental species present, and ecology of the fl data. These features and hydrological meteorological alongside then be considered of climate change must overfi (for example, dam construction, other threats the overall impact. to predict in order or deforestation) useful for all these studies, although Spatial modelling is is often diffi of the results interpretation species. The diffi habitats, and modelling characteristics of underwater of dispersal within catchments, in identifying routes and between and barriers to this dispersal both within catchments. in the The best instrument for evaluating changes standardized ichthyofauna of Africa is long-term, composition, monitoring. This will detect shifts in species 72 CHAPTER 3 | FISH therefore, a key recommendation for theconservation of and management offreshwater andriparianhabitatsis, 1999) (seeChapter 1,thisvolume).Adequate protection (see section3.4)butalsoworldwide (HarrisonandStiassny threats forextinctionof freshwater fi Habitat loss sustainable alternative. biodiversity offers acosteffective and environmentally sustainable managementoffreshwater resources and water securityremain vulnerable.InmuchofAfrica,the in lesswealthynations,where biodiversity the impactsofthreats, butthisinvestmentisnotpossible massive fi of Europe andNorthAmerica)hasonlybeenpossibleby human watersecurityinwealthynations(suchasparts Their studyhasshownthattheprovision ofadequate conservation andfoodsecuritythrough reliable fi freshwater resources goesfarbeyond ensuringbiodiversity have shownthattheneedforsustainablemanagementof recent studies,suchasthatbyVörösmarty can besustainablyexploitedare tothebenefi aquatic ecosystemswithhealthyfi to people.Atthemostbasiclevel,effectively functioning deliver themanyecosystemservicesthatare alsoessential protect andsustainablymanagefreshwater ecosystemsto this iscollectedfortheaquariumtrade–italsoanimportantfoodfi The bandeddistichodus, nancial investmentinwatertechnologytooffset or modifi cation ranksamongtheprimary Distichodus sexfasciatus shes notonlyinAfrica sh populationsthat et al. t ofall.But and (LC), is widespread throughoutcentral Africa.Abeautifulspeciessuchas (LC),iswidespread sheries. human (2010), the more important taxa( as the‘chambo restoration strategicplan’ toprotect oneof end, specialprogrammes havebeen implemented,such the highlevelsofthreat, suchasfrom overfi quite limited,andmore initiativesare required toreduce unique fi especially aimedatprotecting partofLakeMalawi’s Lake MalawiNationalParkwas establishedin1980, al and haveidentifi assessment toidentifypriorityareas for conservation, de laNature (ICCN)initiatedacountry-widebiodiversity Congo andtheInstitutCongolaisPourlaConservation in centralAfricatheMinistryofEnvironment forD.R. conservation ofAfrica’s freshwater fi rivers, lakesandwetlandswillbeimportantforthefuture that specifi or parks.Developmentofparksandprotected areas that fi 2007; Allan borders, where theyare especially vulnerable(Abell freshwater onesthatexistwithinthemoralongtheir many are focusedonterrestrial habitatsratherthanthe exist inAfrica(e.g.,seeStiassny freshwater fi . 2008). sh. shing rightsare stillexertedwithinprotected areas

© SAIAB/ROGERBILLS sh fauna;nevertheless,thescope oftheparkis cally address theconservationchallengesfor shes. Numerous nominalparksandreserves et al . 2010).Forexample,itisnotuncommon ed 30wetlandpriorityareas (Thieme Oreochromis et al . 2011)but,inpractice, shes. Forexample, ) inthelake fi shing. To this shery. et al et . CHAPTER 3 | FISH 73 c sh ciaries of ciaries sheries sheries catch nancial incentives ed by several African ed by several African cation keys in many areas keys in many areas cation shermen and make it available shermen and . 2009). For example, downstream downstream . 2009). For example, et al nancial and logistical means to enforce and logistical means to enforce nancial freshwater ecosystem services pay for the supply of these ecosystem services pay for freshwater services (Forsland for domestic and clean water communities that receive to conserve communities upstream agricultural use pay the to supply clean and manage the habitats so they continue agreements Conservation stewardship and plentiful water. set trust funds are another form of PES scheme, where are up to supply local communities with fi in return (for example, money for new jobs, or healthcare) resources that they manage their freshwater for agreement sustainably. discussed Many of the conservation recommendations by adequate above will be impossible if not backed of regulations development of policy and enforcement governmental many and laws (Smith 2010). However, bodies lack the fi mechanisms must existing laws and rules, and therefore The Convention be put in place to assist in this process. of Biological Diversity (ratifi (Landenbergue countries) and the Ramsar Convention of policy and Peck 2010) may help with the development by the recently and laws. The Convention was supported of the Parties to the at the 10th Conference agreement Convention on Biological Diversity (CBD-COP10, held at and Nagoya in November 2010) that 17% of terrestrial and that globally should be protected, inland water areas species ’By 2020 the extinction of known threatened and their conservation status, has been prevented particularly of those most in decline, has been improved and sustained.’ must be to In conclusion, our objective for the future fi and manage African freshwater conserve effectively at the same time as supporting the livelihoods biodiversity, dependent on these and economies of the people who are the critical stakeholders in ensuring and are resources sustainable management practices. This can only be to scientifi achieved by multi-disciplinary approaches Another recommendation is that reliable fi reliable is that recommendation Another In many available. and made maintained are statistics on species composition information is available little areas, to the lack related quantities. This is directly and catch and identifi of inventories trained personnel. lack of taxonomically and a chronic translate then assimilate and would local staff Trained fi the knowledge of local and scientists to inform decisions managers to resource and policy. Services’ general note, ‘Payment for Ecosystem On a more the benefi mechanisms where are (PES) programs development of tools for the application of research, to conservation and management, and the that research implementation of policy that supports recommendations 2010; made as a consequence of all of the above (Farrell Smith 2010). et . (2009) et al .’s (2007) recommendation (2007) recommendation .’s c programmes, have been c programmes, et al ce (PBWO). Tweddle ce (PBWO). Tweddle c or site-specifi . (2010) discuss the need for a new conceptual . (2010) discuss the need for a new conceptual et al . (2010b) noted that Integrated River Basin Management Allan areas protected framework for developing and managing while that accounts for the need to conserve ecosystems of people. This also allowing for the diverse requirements concept is based on Abell for a multiple-use zoning framework, where focal areas focal areas for a multiple-use zoning framework, where embedded in critical are conservation for freshwater management zones, and these, in turn, embedded in are other programs, catchment management zones. Various management of catchments to large-scale ranging from species-specifi describe some successful projects and conservation describe some successful projects in southern at diverse recommendations Africa directed landscapes to local sites and individual whole scales, from species. These include eradication of invasive species and limitation of the use of alien invasive species in aquaculture actions, at both the site and catchment Such programmes. detail in Chapter 9. in more covered scales, are shown to be important for conservation of freshwater shown to be important for conservation of freshwater of ecosystem services. García biodiversity and provision al to stop (IRBM) is a key conservation action required species decline in northern (2009) described Africa. Cross the catchment-scale actions of the Pangani River Basin by the IUCN- co-ordinated Flow Assessment Initiative (FA), Offi Pangani Basin Water 74 CHAPTER 3 | SPECIES IN THE SPOTLIGHT K The lateral vein creates a vivid red The lateralvein createsavividred for sevenmonthsinanaquarium. melanophores inaspecimenkept Poll (1953)reportedthepresence of pigment (Gerard1936).However, does notresultindevelopment of under lightformorethanonemonth true albino,asplacingliveanimals Heuts 1951)andisconsidereda pigmentation (Boulenger1921; photosensitivity. to theexistenceofextra-ocular typical photonegativereactiondue away fromlight,demonstratinga (1937), notesthatthespeciesmoves contrary toPetit andBesnard 1936). Nevertheless, Thinès(1953), retina andopticalnerve(Gerard lens, andhaveonlyarudimentary embedded inthehead,lacka they arepresent.Theydeeply which literallymeans‘blindbarb’. Kikongo (thelocalNdibu dialect) referred toas‘Nzonzi ampofo’in be discovered.Thespeciesislocally It wasthefi and Stewart1976)ofD. R. Congo. the LowerCongoregion(Roberts from the‘GrottesdeThysville’in on fourspecimenscollectedin1920, Institut Supérieur Pédagogique deMbanza-Ngungu, Bas-Congo, LaboratoiredeBiologie, Democratic RepublicofCongo Royal Museum forCentralAfrica,Vertebrate2 Section, Ichthyology, Leuvensesteenweg 13,B-3080 Tervuren, Belgium 1 A livespecimenof E^le^lËjXcYèfZXm\Ô K_\:fe^fYcè[YXiY1DYXeqX$ The species also lacks The speciesalsolacks Although theeyesarenotvisible, Jg\Z`\jèk_\jgfkc`^_k by Boulenger (1921), based by Boulenger(1921),based was scientifi barb, he enigmatic,Congoblind rst Africancavefi Caecobarbus geertsii Caecobarbus geertsii cally described sh to , from thecave‘GrottedeLukatu’,D.R.Congo. Vreven, E.¹, KimbembimaIbaka,A.²andWamuini Lunkayilakio,S² Kimbembi (2007) and the authors. Kimbembi (2007) andtheauthors. been confi by HeutsandLeleup(1954)has least fouroftheothercavesreported excavation oftheslope. completely disappearedfollowing cave sitein2005foundittohave Leleup 1954).Indeed,avisittothe (Leleup 1956;seealsoHeutsand limestone between1930and1935 extirpated bytheexploitationof One populationwasreportedas de Cristal:750to850melevation). Thysville mountainridge(Monts western slopeandthetopof (formerly Thysville),situatedonthe caves aroundMbanza-Ngungu recorded and HeutsLeleup(1954) several cavesin1949,Heuts(1951) Museum forCentralAfrica. specimen housedattheRoyal total length,basedonthelargest a maximumsizeof80to120mm to beconfi may exceed15years,butthisneeds Romero (2001)statedthelifespan at nineto14years;Proudloveand Heuts (1951)estimatedlongevity abdomen (Petit andBesnard1937). region isvisiblethroughthe a purplishregion,andtheintestinal the operculumgillsarevisibleas band alongthelateralline.Below The presenceof Following explorationsof C. geertsii rmed byrecentsurveys by rmed. The species reaches rmed. Thespeciesreaches from seven fromseven C. geertsii in at inat j_ geertsii River, atributarytotheUombe. The on theUombe orpossibly theKela the Tubulu River andanotherone confi the InkisiRiverbasinseemsto be (2007), thespecies’presencein localities identifi from mappingofthenewcave evidence forthis.However, inferred Inkisi Basin,butatthetimehadno also reportedthespeciesfrom and Daget(1984)Banister(1986) affl (2007) identifi One ofthenewcavesthatKimbembi namely theFuma andtheKokosi. (an affl upper tributariesoftheKwiluBasin C. geertsii Leleup (1954)previouslyconsidered have beenmadeforthese. although nopopulationestimations least smallpopulationsof discovered sevenmorecaveswithat those authors).Kimbembi(2007) (based oninformationsuppliedby would beabout7,000individuals reported byHeutsandLeleup(1954) estimate forthesevencaves 1951); however, agross population and diffi subterranean habitatisextensive have beenimpossiblebecausethe Statistical populationsurveys © ROYAL MUSEUMFORCENTRALAFRICA Heuts (1951) and Heuts and Heuts (1951)and uent of the Kwilu Basin. Lévêque uent oftheKwiluBasin. Lévêque rmed by two of them – one on rmed bytwoofthem –oneon uent of the Lower Congo), uent oftheLowerCongo), isontheTobo River, another cult to sample (Heuts cult tosample(Heuts to be present in only two tobepresentinonlytwo ed asholding ed by Kimbembi ed byKimbembi C. geertsii C. C. , CHAPTER 3 presence of C. geertsii in D. R. Congo, of the caves inhabited by C. 1,500 individuals to the Unites as reported by Lévêque and Daget geertsii, other typical cave animals, States (Proudlove and Romero (1984), is incorrect. Thus, the entire such as terrestrial insects, are 2001). Three other primary threats distribution area of the species is absent. Therefore, C. geertsii is to the species were identifi ed by about 120km2. Heuts (1951) noted entirely dependent on an external, Brown and Abell (2005): changes | important differences between the exogenous, food supply to the caves in hydrology of the small rivers SPECIES IN THE SPOTLIGHT different populations of C. geertsii during the rainy season with, as a feeding the caves; increasing in the Kwilu basin. Populations result, important fl uctuations in human population; and associated present in affl uents of the Kokosi food resources between seasons. deforestation (Kamdem Toham River have an opercular guanine Moelants (2009) states that et al. 2006). Since 2003, with the spot which may cover one third of the species may feed on small attenuation of the political situation the operculum (in addition to a few crustaceans living in the caves, in D. R. Congo and the rehabilitation other guanine spots and marks). but this needs to be confi rmed. of the Matadi-Kinshasa road, there This spot is absent in all other Consequently, growth is extremely has been a signifi cant infl ux of populations (affl uents of the Fuma slow, and all further available data rural people towards Mbanza- River). Furthermore, within one suggest a very low reproduction rate, Ngungu. Consequently, land use has

cave the population has a serrated justifying protection measurements. increased around Mbanza-Ngungu dorsal spine, which was not found in A visit to the Kambu cave by the for buildings as well as agriculture. all other populations examined by authors in August 2009 failed to fi nd One cave is now used as a quarry, Heuts (1951). the species, although its presence with consequential loss of the Traditionally, caves are sacred had been reported by Kimbembi Caecobarbus population (Leleup in the area (Laman 1962) and, as a (2007). However, several individuals 1956; Poll 1956; and see above), and result, access to most of the caves is of at least one species of Clarias (± others are at risk of collapse due restricted still today. A law, passed 200mm standard length) were found to human disturbance (Kimbembi on 21 April 1937, protected C. geertsii in the different isolated pools. This 2007; Moelants 2009). Agriculture is from all hunting and fi shing, except observation suggests predation of practiced preferentially in the valleys for scientifi c purposes (Frenchkop C. geertsii by species of Clarias, as near to the caves but may also occur 1941, 1947, 1953; Duren 1943). The previously proposed by Heuts and on the hillside slopes surrounding species was added to the CITES Leleup (1954) and by Leleup (1956). and covering the caves, leading to Annex II (on 6 June 1981), resulting Caecobarbus geertsii has, in the increased erosion and landslides. In in an international trade restriction past, been traded as an aquarium the past, these areas were covered which means that the species cannot fi sh, with large numbers having with lowland rainforest and be traded without appropriate been exported to industrialized secondary grassland (White 1986), export and import permits. C. geertsii nations. Collection pressure should limiting erosion. Further research is still the only African freshwater have been reduced through listing and conservation initiatives in the fi sh species on the CITES list. The under CITES; however, a CITES fi eld are necessary if this unique IUCN Red List status of C. geertsii certifi cate was issued to import species of fi sh is to survive. is Vulnerable (VU), due to a limited geographic range and a decline in the area and quality of its habitat (Moelants 2009). Caecobarbus geertsii was found in only seven of the 45 caves explored by Heuts and Leleup in 1949. This indicates, according to Heuts (1951) and Heuts and Leleup (1954), that caves must have a specifi c combination of ecological conditions if they are to be populated by C. geertsii, and they summarised the following conditions: 1. high calcium bicarbonate concentrations in the water; and 2. a distinct periodicity of the subterranean river fl ow regime Land use around the entrance of the ‘Grotte de Lukatu’, with subsequent landslides through the caves. visible (9 March 2007). The entrance to the cave is directly below the largest trees Due to this periodic inundation in the middle of the photograph. © ROYAL MUSEUM FOR CENTRAL AFRICA 75 76 CHAPTER 3 | SPECIES IN THE SPOTLIGHT K rivers (Abban fi value and,consequently, thehigh indicated bytheirhighmarket popularity oftilapiainAfricais to othernativefi recognised asasignifi niloticus famous Nile tilapia( in manypartsoftheworld,and common sourcesofinvasivespecies Aquaculture isalsooneofthemost areas outsideoftheirnativerange. been introduced,orescaped,to become aproblemwheretheyhave over othertilapia,whichcan niloticus 1990). Allofthesefactorsaccord and highfecundity(Ogutu-Ohwayo growth rate;largemaximumsize; Khallaf andAln-Na-Ei 1987);high and Moriarty1973;Getachew1987; levels (Moriarty1973;Moriarty food spectrumfromvarioustrophic a widevarietyofhabitats;its its abilitytoestablishandoccupy attributed toseveralcharacteristics: fi Africa. Itssuccessasacommercially InstitutdeRecherche pourleDéveloppement, 213rueLaFayette 75180Paris Cedex10,France 3 DépartementBiologie Intégrative,CNRS UMR5554,Université deMontpellier IICC 63PlaceEugèneBataillon F-34095Montpel 2 National Museums ofKenya,P.O. Box40658Nairobi, 00100,Kenya 1 Trewavas, 1983 intheSuguta niloticus six sub-speciesofNile tilapia: Eastern Africaisendowedwith K_\E`c\k`cXg`X et al K`cXg`Xè\Xjk\ie8]i`ZXÆX]i`\e[Xe[]f\ elsewhere; Basin butnowwidelyintroduced originally fromtheWhite Nile Lake Turkana; niloticus vulcani Kivu, AlbertandGeorge; (Boulenger, 1912)inLakesEdward, shing pressureinmostlakesand shed andculturedspeciesis . 1994). Jg\Z`\jèk_\jgfkc`^_k to somanypeopleacross industry thatisimportant much oftheaquaculture ilapia formthebasisfor with greatcompetitiveness O. niloticuseduardianus et al. O. niloticus sugutae (Trewavas, 1933) in sh species.The ), inparticular, is

(Linnaeus, 1758), 2004;Gréboval Oreochromis cant threat O. O. O. O.

O. O.

O. esculentus stocks ofnativetilapiaspecies 1954 and1962,duetodecreasing ﬁ the purposeofimprovingtilapia introduced toLakeVictoria for variabilis two nativetilapiaspecies( caused thedisappearanceof the introductionof Twongo 1995).Itisthoughtthat (Mann 1970;Ogutu-Ohwayo1990; established ininshorehabitats the endof1960swaswell colonized theentirelakeandby Oreochromis niloticus (Nyingi and oneotherrecentlydiscovered Trewavas, 1983inLakeBaringo; river basin; known foritsabilitytohybridize esculentus the mainpartoflake– esculentus the declineof O. niloticus hybridization withsubspecies of It wasinitiallyhypothesisedthat the bulkofﬁ Rift Valley. Baringo andBogoriaintheKenyan the Loboiswamp,betweenLake Lake BogoriaHotelspringnear undescribed subspeciesfromthe aquaculture. aquaculture. speciesfor (LC), ahighlyfavoured The Niletilapia, sheries inseveralphasesbetween Oreochromis niloticus et al. and , because havingoncerepresented wasthemaindriverof O. niloticusbaringoensis © LUCDEVOS and 2009), butstill O. esculentus O. variabilis Oreochromis niloticus sheries inthelake. O. variabilis. O. niloticus O. niloticus rapidly

was ) from and O. O. O. O. is well O. O.

, Nyingi, D.W.¹ andAgnèse,J.-F.² O. niloticus the competitivesuperiorityof Nyingi andAgnèse Rognon andGuyomard2003; 1988; MwanjaandKaufman1995; with othertilapiines(Welcomme provision ofsuitablespeciesfor is stillneededtoensurethe advances, considerableinvestment of suitablefeed.Despitethese and inresearchproduction provision ofseedfi the centrehasbeeneffi water freshwaterspecies.Sofar, provides fi under theFisheriesDepartment, In Kenya,theSaganaFishfarm, shillings (EUR70,000)forponds. geared toreceive8millionKenya existing plans,eachconstituencyis by June2013.Accordingto fi Development toconstruct200 by theMinistryofFisheries (EUR 10.67million)waslaunched worth 1,120millionKenyashillings the fi farmers andotherstakeholdersin country byraisingtheincomeof a viableeconomicactivityinthe been tohighlightfi intentionhas The government’s Economic StimulusProgramme. to promoteaquacultureunderthe government hassteppedupefforts foreign fi sector initiatedanddependenton with allnewactivitiesinthe eastern Africahasbeenfi development ofaquaculturein The greatestlimitationto K`cXg`XXe[XhlXZlckli\ (Balirwa 1992;Agnèse contribution fortheirextinction demonstrated tobethemostlikely two formernativespecieswas sh pondsin140constituencies shing industry. Theproject, nancing. InKenya,the ngerlings forwarm

subspecies lier Cedex5,France sh tofarmers sh farmingas

2007). However,

over the et al. cient in nancial, 1999). , ³ CHAPTER 3 the various regions, ensuring when it was introduced to Lake baringoensis. Even though impacts development of the industry. Victoria. Unfortunately, O. niloticus of the possible introduction of With the government supporting has, in many cases, been the species O. leucosticus are still unknown, new initiatives, the greatest of choice for aquaculture, therefore introductions of tilapiines continue challenge is to identify a suitable leading to further problems of to be made within the region, | species that will ensure high competition and hybridisation. either intentionally or accidentally SPECIES IN THE SPOTLIGHT yield, while also safeguarding Oreochromis leucosticus was through escape from culture ponds. native species from the impacts of originally known from drainages Such issues are a clear indication introduced aquaculture species. near the border of Uganda and of a failure of well-defi ned policies, Unfortunately, in Africa the search the D. R. Congo, specifi cally or implementation of the existing for suitable species for aquaculture Lakes Edward, and Albert, and regulations, for the management has often disregarded potential associated affl uents. However, it of natural fi sheries resources in impacts on the native species. was introduced to Lake Naivasha Kenya. Through lack of awareness, The most important culture in Kenya in 1957 (Harper et al. and desperation to increase species are still mainly taken 1990). About 150km away from yield, fi sh farmers are breeding from the wild, and populations Lake Naivasha is Lake Baringo, in alien species of tilapia that could

are often translocated to basins the Kenyan Rift Valley, home to the naturally hybridize in a similar far beyond their native range, endemic subspecies of Nile tilapia, manner – as seems to have occurred potentially bringing closely related O. niloticus baringoensis. Nyingi in Lake Baringo. Consequently, but formerly isolated species or and Agnèse (2007) note that O. native species may be lost in several populations into contact with niloticus baringoensis share genetic parts of eastern Africa, as already each other. Where there has been characteristics of O. leucosticus, observed in Lake Victoria. inadequate research and planning, suggesting that O. leucosticus might As noted above, a new sub- an introduced cultured species have been introduced also to Lake species of Oreochromis was recently may directly compete with native Baringo, with some subsequent discovered from the Lake Bogoria species, or may hybridize with transfer of genetic material through Hotel spring near the Loboi swamp. them, as noted above for O. niloticus hybridization with O. niloticus This population was formerly

Fish farms, such as this one in Malawi, represent an important source of food and income for people throughout Africa. However the traits that make species such as Oreochromis niloticus suitable for aquaculture mean that they pose a signifi cant threat to local species should they escape. © RANDALL BRUMMETT 77 78 CHAPTER 3 | SPECIES IN THE SPOTLIGHT Baringo andpartlytoLake Bogoria. River nowpartly fl Loboi andSandai Rivers.TheSandai caused changesinthecourses of the induced heavyrainsof1997,which recent wasduringtheElNiño- of riversinthisregion.Themost have causedchangesinthecourse In addition,periodicavulsions (Ashley from macrophytesintheswamp aerobic decompositionofdetritus of highoxygenconsumptionduring discharges), whichisaconsequence in thespringsandgroundwater oxygen, comparedtoaround60% level (around4%saturateddissolved signifi to LakeBaringo.Theswamphasa Loboi River, whichdrainsfromit fl new sub-speciesisfound)that warm watersprings(wherethe and chemicalbarrierbetweenthe the Loboiswampactsasaphysical and Agnèse2007).Themainbodyof originality (Nyingi 2007;Nyingi analysis demonstratedits but geneticandmorphological thought tohavebeenintroduced, (Ashley abstraction forirrigationsince1970 over thelast30yearsduetowater itself hasrecededbyaround60% encroachment. TheLoboiswamp rivers isunderthreatfromhuman the Loboiswampandassociated determination in may beamodelforthestudyofsex Therefore, thenewsub-species D’Cotta 2001;Tessema high temperatures(Baroillerand is knowntobeinfl sex-ratio, sincesexdetermination special mechanismstoregulateits sub-species mayalsohavedeveloped oxygen levelsmayalsobelow. This resistance mechanismsasdissolved and mayhavedevelopedhypoxic temperatures (approximately36°C) The sub-speciesinhabitshigh development, ifmanagedproperly. new possibilitiesforaquaculture Loboi swampoffersinteresting from thespringsdraininginto ow intotheswamp,and However, thepopulationfrom Thenewapparentsub-species cantly lowdissolvedoxygen et al. et al.

2004; Owen 2004). Oreochromis. uenced by ows intoLake et al. et al. 2006).

2004). al. of theriverbanks(Harper local farmersleadingtoweakening agricultural encroachmentby of fl to LakeBogoria.Thesechanges changed itscourseandnowfl used tofeedLakeBaringo,has Similarly, theLoboiRiver, which Ô DXeX^\d\ekf]k`cXg`X to manage thelakeasa complete exploitation have madeitdiffi regulations and prioritiesfor (Marten 1979).Thesedifferent stocks targetedforexploitation fi when theyadopteddifferent (Kenya, Uganda andTanzania), countries borderingthelake following independenceofthe compromised intheearly1960s of LakeVictoria washighly example, fi shared bydifferentcountries.For where freshwaterresourcesare A signifi replicate thesesuccesseselsewhere. Kenya. Thereis,however, aneedto protect hydrologicalsystemsin Management Authorityisableto that theNational Environment of thisprojectisaclearindication Nairobi (Nzioka 2009).Thesuccess within thecityandsuburbsof washes, andotherhumanactivities disposal, sewage,run-offfromcar greatly impactedduetosolidwaste Nairobi River, whichhadbeen ensuring rehabilitationofthe has beenactivelyinvolvedin Management AuthorityinKenya Kenya. TheNational Environment almost alllakesandriversystemsin Loboi swampbutiscommonin situation isnotuniquetothe and education. programmes ofpublicawareness management actionwith to reinforce policyand shing regulationsbasedonthe j_\i`\j

2003; Owen ow werealsoduetointensive cant challengehasexisted sheries management ...many challenges still lie ...many challengesstilllie ahead, anditwillbecritical et al.

2004). This et ows cult ecosystem (Ntiba awareness andeducation. action withprogrammes ofpublic reinforce policy andmanagement lie ahead,anditwillbecritical to Nevertheless, manychallengesstill (LVFO 2007;Lubovich2009). a harmonizedfi their nationalpoliciesanddevelop respective governmentstoreview Activity; thisplancalledonthe for theManagementofFishing adopted aRegionalPlanofAction 2007, Kenya,Tanzania, andUganda resources. For example,inMarch management ofthelake’sfi programs forsustainable consistent setofpoliciesand of providingacomprehensive, being madetoaddressthechallenge conducted. Signifi which fi highly explicitonthemannerin Fisheries Actof1991,whichis weakness inenforcementofthe possible becauseofthelaxityand this overexploitationhasbeen to overexploitation.Inthepast, 2009), butisalsoprobablydue catches (Njiru individuals observedincommercial et al. allowed totallengthof30cm(Njiru fi were dominatedbyimmature to 2005showthatartisanalcatches surveys of1998to2000and2004 perch ( numbers ofintroducedNile be partlyduetotheincreased Stock analysesfor native sub-species(seeabove). responsible forthedeclinein niloticus to declinesintheintroduced of managementhascontributed et al. sh, mostbeingbelowthelegally 2007).Thepaucityofmature 2005).Ironically, thislack Lates niloticus shing activitiesshouldbe , whichwaspreviously et al shing framework

cant effortsare et al. O. niloticus . 2005)may ) (Lubovich 2001; Njiru shery

O. O.

CHAPTER 3 Jg\Z`\j`ek_\jgfkc`^_k =fi\jki\deXekj`en\jk\ie8]i`ZXÆ |

mXe`j_`e^`jcXe[jf]jpcmXeÔj_\j SPECIES IN THE SPOTLIGHT

Moritz, T. ¹

signiﬁ cant part of western Africa is covered by differing types of savanna that are drained by a few 8large rivers, like the Niger, Volta and the Senegal. The vegetation reﬂ ects climatic conditions

including a cycle of dry and wet seasons. Closer to the coast, partly bordered by the Guinean highlands (from the highlands of the southern Fouta Djallon in south-eastern Guinea, through northern Sierra Leone and Liberia, to north- western Côte d’Ivoire), the climate is more humid, allowing different types of forest to grow. These forests are inhabited by animals Lokoli swamp forest in southern Benin. This small forest fragment serves as one of closely resembling or even identical the last refuges for many forest dwelling species. © T. MORITZ to those of the central African forests. Thus, many sylvan (forest dwelling) fi sh species and species- species. Examples include the Cfbfc`]fi\jkÆXi\]l^\ groups fi nd their most westerly fourspine leaffi sh (Afrononandus An exemplary forest remnant distributions within the western sheljuzhkoi) and the African leaffi sh is the Lokoli swamp forest in Africa coastal forests. A number of (Polycentropsis abbreviata), with southern Benin. This small, these westerly sub-populations may their closest relatives in Asia and (approximately 500ha) piece of be discrete sub-species, or separate South America, and the enigmatic forest is permanently fl ooded by a species within species complexes, denticle herring (Denticeps network of channels from the Hlan showing distinct colour morphs, or clupeoides), which is the only River, an affl uent of the Ouémé other unique features. extant representative of the family River. It is approximately 20km The high number of unconnected Denticipetidae, the sister group of east of Bohicon and 100km north coastal rivers in western Africa is all other clupeomorphs. of Cotonou and can only be crossed thought to have promoted these The climatic conditions of the by boat. The forest is densely speciation processes, which have Guinean region not only provide vegetated with high tree density, led to noticeably high levels of good conditions for forest and the tree cover is usually closed endemism, such as for characids, ecosystems, but also support a more above the channels. Most channels barbs and cichlids. Furthermore, diverse and reliable agriculture are less than a metre wide and are several remarkable fi shes, which compared with the Sahelo-Sudan only navigable using small dugout are sometimes called ‘relict‘ species, region. This promotes better canoes. Water depth varies by less occur in the Guinean regions. livelihood opportunities which, in than one metre within a year, and is These species belong to phyletically turn, lead to increased population usually around 1 to 2.5 metres. The old groups previously represented densities and a greater demand for water has a dark brown colouration by more numerous and widespread land. With increased demands for due to leaf litter decomposition, a species but, following evolutionary agricultural land, deforestation moderate acidic pH of 6 to 7 and a events, now represented by only continues, leaving only forest temperature of around 26°C. The a few, often locally restricted fragments in some areas. channel substrate is predominantly

1 Deutsches Meeresmuseum, Museum für Meereskunde und Fischerei – Aquarium, Stralsund, Germany 79 80 CHAPTER 3 | SPECIES IN THE SPOTLIGHT al are atthe mostwestern pointsof ( buchholzi calabaricus coast. Thereedfi network oflagoonsparallelto the the Niger Deltaandtheconnected from thecoastalforestedrivers of forest species,otherwiseknown Lokoli are,toahighdegree,typical of theOuéméRiver, fi connection tothemainchannel is verydifferent.Despiteadirect (Rödel species ofreptilesandamphibians relatively fewexclusiveforest herpetological surveyshaveshown which isendemictoBenin.While ( and thered-belliedmonkey including pangolins,fl animals intheDahomeyGap, the lastrefugesforforestdwelling layer ofvariabledepth. places haveamudandleaflitter where thecurrentisstronger;most sand, withsmallpatchesofgravel © T. MORITZ Barboides britzi Gnathonemus petersii Cercopithecus erythrogaster . 2010),thesituationforfi The Lokoliforestservesasoneof et al. ), andelephant nose fi ), butterfl 2007; Ullenbruch , oneofAfrica’s fi smallestfreshwater sh ( y fi Erpetoichthys ) inLokoli shes ofthe ying squirrels sh ( ), Pantodon shes et sh 2007; pers.obs.).Thecyprinidgenus their ranges(Montchowui Barboides source of incomeandbush meat meat traderemains animportant regulations for hunting, thebush taboos prescribingatleastsome the forestitself,despitereligious along theforestmargins.Within extensive clearanceforagriculture fragment aresignifi (Moritz 2009). small affl was onlyrecentlydescribed,froma distichodontid example, thebottomdwelling in suchunusualhabitats.For smaller bodysize,awaitdiscovery more fi and Moritz2006).Itislikelythat 12.6mm standardlength(Conway other freshwaterfi mature atasmallersizethanany miniature fi to theLokoliforestitself.This of itsrange,with is alsoatthemostwesterlypoint smallest Africanfreshwaterspecies, Impacts tothissmallforest sh species,especiallyof inBenin. sh, isanewlydescribedspeciesendemictoLokoliforest uent oftheLokoliforest , consistingoftwothe sh becomessexually Nannocharax signifer B. britzi sh inAfrica,at cant, with endemic et al.

palm by cuttingoffthetopsof secondary productsareproduced leading toCotonou.Palm wineand is openlysoldalongthemainroad as thetaro( introduction ofalienspeciessuch has beenfurtherimpactedthrough over-harvesting. Theforestfl signifi formerly dominantpalmhasbeen As aresult,theabundanceofthis impacts totheplantsthemselves. river isclearly visibleon aerialor The courseofthis smallcoastal Nigeria provides agoodexample. River attheborderofBeninand expanding agriculture.TheIguidi more, habitatdegradationdue to areas, theprimaryproblemis, once which isheavilyimpactingcertain threats. Inadditiontopollution, Guinean forestedrivers,facesimilar more spaciousthansomeofthe forest. even withinclearingsintheswamp tubers, whichiswidelyplanted, introduced plantvaluedforitsroot Forested coastalrivers,although Raphia hookeri, cantly reducedthrough Colocasia esculenta withevident ora ), an CHAPTER 3 satellite images, due to its bordering the Lokoli, can sustain a number distributions, they may be of high gallery forest. The forest stands out of small endemic species. What importance for maintaining genetic in stark contrast to neighbouring, is more important, however, is variability within a species and in and continuously expanding, fi elds. the complexity of biodiversity ongoing evolutionary processes. The Iguidi River fl ows in a north- that is found and that needs to be Therefore, despite their small size, | south direction, starting out as a conserved. Forest fragments and forest fragments deserve greater SPECIES IN THE SPOTLIGHT small forested stream that develops remnants of gallery forests are focal focus within conservation plans; into a swamp. As is typical for a points of habitat complexity, edge their inclusion will help to ensure forest stream, the water is brown to effects and ecological interactions preservation of biodiversity in all dark brown in colour, although the – and as outposts for species its forms. pH is not especially acidic, at 6.5 to 7.5; water temperature is commonly 26 to 29°C; and conductivity is low at 50 to 65μS (Moritz 2010). Despite the river’s low salt content, fi shes characteristic of brackish

environments are also present, such as the freshwater pipefi sh of the genus Enneacampus, and the sleeper goby (Eleotris daganensis). The majority of fi shes from the Iguidi are, however, typically freshwater, forest-dwelling species such as the dotted catfi sh (Parauchenoglanis monkei), the small distichodontid, Neolebias ansorgii, and the cryptic mormyrid (Isichthys henryi). This small river represents an outpost of the Lower Guinean forest, and holds the most westerly distributions of several Lower Guinean species, such as the aforementioned Neolebias ansorgii, the Niger tetra (Arnoldichthys spilopterus), and the catfi sh Schilbe brevianalis (Moritz 2010). Furthermore, the Iguidi River is the type locality for the rare, miniature Barbus sylvaticus, the even smaller Barboides gracilis, and the denticle herring (Denticeps clupeoides), all of which are assessed as Vulnerable or Endangered. In conclusion, at fi rst glance, small forest fragments seem to be of minor importance for the conservation of forest dwelling species – often being too small to sustain endemic species, or even too small to harbour a discrete population of a sylvan species. Many inhabitants of forest remnants are, therefore, The freshwater butterfl yfi sh, Pantodon buchholzi (LC), a widespread species in non-specialist or even savanna Africa, reaches the most westerly point of its range in Lokoli forest, Benin. This species. A closer view of the fi shes, species is capable of jumping out of the water to search for insects or to escape however, reveals a quite different from predators. It is not a glider, but a ballistic jumper, with tremendous jumping picture. Forest remnants, such as power. © T. MORITZ 81 82 CHAPTER 3 | SPECIES IN THE SPOTLIGHT C (Nile tilapia)and grounds for the eastoflakeserveasbreeding Tana ecosystem.Thewetlandsto are anintegralpartofthecomplex largest wetlandareainEthiopiaand surround mostofthelakeform Khartoum, Sudan. total volumeoftheNile Riverat contributing morethan80%ofthe only riverfl headwaters oftheBlueNile –the Lake Tana, whichitselfformsthe Today, severalriversdraininto fi which areimportantforthelake (North Africancatfi (Mohr 1962;Chorowicz of theBlueNile nearitssource long basaltfl early Pleistocenewhena50km- 1,800m. Itwasformedduringthe at analtitudeofapproximately in thenorth-westernhighlands largest inEthiopia.Itissituated k_\ 8le`hl\jg\Z`\jÕ Department of Biology, AddisAbaba University, Ethiopia 1 as aseparatesub-species, niloticus a surfaceareaof3,150km Labeobarbus. belonging tothecyprinidgenus Nemacheilus. 2007), group’; seeDeWeirdt andTeugels barbs’), Labeobarbus the genera fauna includesrepresentativesof (Vijverberg to thelakeanditscatchments Lake Tana, ofwhich20areendemic sheries (Vijverberg The wetlandsandfl There are28speciesoffi Garra, Varicorhinus endemic speciesfl it, arehometoaunique, the riversthatdraininto ake Tana, inEthiopia,and Barbus inLakeTana wasdescribed Jg\Z`\jèk_\jgfkc`^_k CXY\fYXiYlj Oreochromis, Clarias, owing outofthelake, et al. Oreochromis niloticus (i.e., the ‘large African (i.e., the‘largeAfrican ow blockedthecourse

The populationof The lake,whichhas (i.e., the‘small 2009). Theﬁ Clarias gariepinus sh), bothof et al oodplains that . 2009). . 2009). ock Oreochromis et al. and and 2 sh in , isthe sh sh Barbus 1998). O. O.

Gambusia holbrooki niloticus tana Garra, Labeobarbus represented byfourgenera, species-rich familyinthelake, The Cyprinidaearethemost K_\ recent times. trace ofthesefi Meskel 1981);thereis,however, no introduced intothelake(Tedla and from Italyduringthelate1930sand were reportedtohavebeenbrought Within the out completely as recently as 16,000 out completely as recently16,000 evidence suggeststhelakedried for somanynewspecies,ashistorical interesting isthespeedofevolution from theoutfl Falls, located30kmdownstream remained isolatedduetotheTissisat colonising species,andithassince radiation amongtheoriginal that mayhavepromotedadaptive it offeredseveralnewhabitats species because,whenitfi a largenumberofcloselyrelated that thelakeisabletosupportsuch and Dejeninprep.).Itisthought complex of17species(Getahun component oftheLakeTana fi Labeobarbus macrophtalmus CXY\fYXiYlj Zfdgc\o fZbèCXb\KXeXÆ Labeobarbus . Two exoticspecies, ow of the lake. Most ow ofthelake.Most shes from the lake in shes fromthelakein

and and jg\Z`\jÕ

Varicorhinus. is a unique isaunique Esox lucius rst formed, is a benthopelagic species that forms animportant isabenthopelagicspeciesthatforms Barbus shery. fZb , , © LEONAGELKERKE

fewer than15,000years(Vijverberg species complexmayhavetaken the evolutionof years ago(Lamb categories of spawning behaviour are categories ofspawning behaviourare dissolved oxygencontent.Three gravel beds,andaslightlyhigher which hasmoretributarieswith numbers migrateuptheMegech, 2005), althoughslightlygreater Dirma riverstospawn(Anteneh Lake Tana upboththeMegechand species werefoundtomigratefrom and easternpartsofthelake.Five for thesespeciesinthenorthern provide idealbreedinggrounds Dirma Riversandtheirtributaries indicate thattheRibb,Megechand Limited surveysaroundLakeTana JgXne`e^Y\_Xm`fli et al dominant species offshore. dominant speciesoffshore. tsanensis species attherivermouths. habitats andarethepredominant tsanensis for spawning. periodically migrateintotherivers are piscivores,andmostofthem Eight ofthe . 2009). . 2009). and are abundant in the inshore are abundantintheinshore

L. brevicephalus L. intermedius Labeobarbus et al Labeobarbus . 2007), meaning . 2007),meaning Getahun, A.¹ species species and L. L. are the arethe L. L.

CHAPTER 3 observed (Anteneh 2005), obligate employs gillnets, and the main target The same authors have reported a river spawners, lake spawners and species is Nile tilapia (O. niloticus). 75% decline (in biomass) and 80% generalists (spawning in both the However, the reed boat (tankwa) (in number) of landed fi sh of the lake and its tributary rivers). fi shermen also use hooks and lines, species of Labeobarbus (L. acutirostris, At least seven species spawn in and traps, as well as spears to catch L. brevicephalus, L. intermedius, L. | the headwaters of the main rivers catfi sh. macrophthalmus, L. platydorsus and SPECIES IN THE SPOTLIGHT draining to the lake. As yet, there is In 1986, motorised boats and L. tsanensis) in the southern gulf no evidence of river-specifi city, but nylon gill nets were introduced of Lake Tana. The most plausible this cannot be discounted. After a as part of the Lake Tana Fisheries explanation for the decline is brief pre-spawning aggregation at Resource Development Program recruitment overfi shing by the the river mouths, the adults migrate (LTFRDP) (Anteneh 2005). Data commercial gillnet fi shery (de Graff upstream in July and August, at collected from all commercial et al. 2004), and poisoning of the the onset of the rainy season. Final fi sheries recognizes only four spawning stock in rivers using the maturation and spawning occur in species groups: Labeobarbus spp., crushed seeds of ‘birbira’ (Milletia the tributaries of the major rivers, or African catfi sh (C. gariepinus), ferruginea) (Nagelkerke and Sibbing possibly in gravel reaches in the main Nile tilapia (O. niloticus) and beso 1996; Ameha 2004).

channels. After spawning, the adults (Varocorhinus beso). This fi shery The commercial gill net fi shery return to the lake for feeding until mainly supplies larger markets, for species of Labeobarbus is the next cycle of breeding. Highly using 100m long gillnets. There are highly seasonal and mainly targets oxygenated water and gravel beds are around 25 motorised fi shing boats, spawning aggregations, as more important for development of the most of which land their catch in than 50% of the annual catch is eggs and larvae. Deposition of eggs Bahir Dar, the main town on the obtained in the river mouths during in gravel beds prevents them from shore of Lake Tana. The fi shery August and September. There is being washed away, and clear water is, however, expanding to all 10 also a chase and trap fi shery based is required to ensure they are free of Woredas (districts) bordering the in the southern part of the lake, and sediments that might obstruct the lake, including the Gorgora area (on longlines, cast nets and traps are diffusion of oxygen. the northern shore). occasionally used but contribute The juveniles start to return to Total annual catches increased little to the total fi sh catch. the lake in September and October from 39 tonnes in 1987 to 360 tonnes as fl ows reduce, where they feed in 1997 (Wudneh 1998). However, K_i\XkjkfCXb\KXeXXe[`kj and grow to sexual maturity. There the catch per unit effort for the CXY\fYXiYljjg\Z`\j is good evidence that, during their commercial gill net fi shery targeting Overfi shing return to the lake, the juveniles may Labeobarbus species dropped by Although a fi shery policy has been remain in the pools of the main river more than 50% over the period developed both at federal and segments for an extended period, 1991 to 2001 (de Graff et al. 2004). regional levels, it is not effectively probably until the next rainy season, at which time they will be carried into the lake.

K_\cXb\Ôj_\i`\j The lake fi shery is clearly very important to the local population, employing more than 3,000 people in fi shing, marketing, and processing (Anteneh 2005). Traditionally, the main fi shery has been a subsistence reed boat fi shery targeting a range of species, sometimes including the Labeobarbus species. This was conducted throughout the lake until the 1980s; since then it has been replaced in many areas by other methods. The fi shery remains important in the more remote areas of the lake, with the catch being Fishermen cast their lines from papyrus boats ("tankwas") on Lake Tana in northern sold at small markets or used for Ethiopia. Behind them lies the source of the Blue Nile. Near Bahir Dar, Ethiopia. household consumption. It mainly © A. DAVEY 83 84 CHAPTER 3 | SPECIES IN THE SPOTLIGHT Bahir Dar. the southaroundtownof some partsofthelake,especiallyin chemicals, suchas‘birbira’ ( to waterquality inthelake.Other their usepresents apotentialthreat a lackofeffectiveregulationon is stillrelativelylimited;however, agricultural productsbyfarmers into thelake.Theuseofthese (including DDT),andherbicides agricultural fertilizers,pesticides around thelakeisbringing Run-off fromsmall-scaleagriculture Water pollution excavation ofsand. land isnowusedforcultivationand around shores.Someoftheexposed available aquatichabitat,especially has resultedinfragmentationofthe This reductioninthewaterlevel and alossofwaterholdingcapacity. lake’s area,adropinwaterlevels, lake haveledtoareductioninthe huge depositsofsedimentintothe et al. tonnes perhectareyear(Teshale around thelakeisbetween31and50 areas. Thesoillossratefromareas and smotheringupstreamspawning sediments drainingintothelake for soilerosion,resultingin widespread, facilitatingconditions deforestation hasbecomevery macrophyte standsinotherareas. there isanexpansionofsubmerged harvesting andburning,while emergent macrophytesthrough with thesubsequentdepletionof wetlands increaseseveryyear, Human encroachmentonthe ‘fl people usetheshoresoflakefor As seasonalfl Habitat disturbance led tooverfi gears. Asreportedabove,thishas there islittleregulationoffi exploiting thefi illegal, unregisteredfi to everyone.Therearestillmany resources thatarefreelyavailable are, unoffi implemented. Lakesandrivers oodplain recession agriculture’. oodplain recessionagriculture’. Over the last 15 years, Over thelast15years, 2001; Teshale 2003).These cially, consideredtobe shing of ooding recedes,many sh resources, and sh resources,and Labeobarbus shermen shermen Milletia shing shing in

Tana and itstributaries, with the waterlevel and qualityinLake purposes. These projectsmaylower directly fromthelakeforirrigation River; andsometopumpwater discharging thewaterinto Beles to ahydropowerfacilitybefore through tunnelsfromthelake store water;sometopumpwater to impoundthelake’stributaries projects. Someoftheseareintended and Gumaradamsirrigation Koga, Ribb,Megech,GilgelAbay basin watertransferproject,andthe These includetheTana Belesinter- consideration orbeingimplemented. dam andirrigationprojectsunder governments, thereareseveralother by thefederalandregional is consideredagrowthcorridor Lake Tana andBelessub-catchment projects. However, because the privately run,small-scaleirrigation points aroundthelakeasaresultof Water abstractionoccursatsome impoundment Water abstractionand fauna, including pollute thelakeandkillaquatic ichthyocide; seeabove),mayalso ferruginia mitigate thesepollutionthreats. sewage systemcouldsolveor the developmentofanappropriate discharged directlyintoLakeTana – town ofBahirDaris,inmostcases, Domestic wastewaterfromthe spawn in gravel beds, such as seen here intheGumaraRiver.spawn ingravelbeds,suchasseenhere Many ofthe ) seed powder (used as an ) seedpowder(usedasan Labeobarbus Labeobarbus species migrateuptheriversfl species. species. and implemented. suggested willbestrictlyfollowed and themanagementplans recommended mitigationmeasures projects, soitishopedthatthe been conductedformanyofthese assessment (EIA)studieshave cyprinids. Environmentalimpact extinction ofthisuniquefl blocked, potentiallyleadingtothe newly constructeddams,maybe measures toallowpassagepast migrations that,withouteffective of As reportedabove,manyspecies subsequent impactstobiodiversity. future. this lakeistobe conservedforthe required iftheunique fi action byallstakeholdersis solving thisproblem.Concerted basin authorityisanoptionfor development ofaLakeTana sub- any recommendationsmade.The capacity toenforceandimplement EIAs stilllackthestrengthand the implementingagenciesfor with implementation.Inaddition, the EIAstudiesandfollowup mitigation measuresinsomeof makes itdiffi Labeobarbus ecology ofthedifferentspecies on thebiology, behaviour, and Comprehensive scientifi institutional capacity Lack ofinformationand Labeobarbus owing intoLakeTana to are stilllacking.This cult torecommend undergospawning © LEONAGELKERKE

sh faunaof c studies ock of CHAPTER 3 The Twee redfi n, Barbus erubescens, a Critically Endangered fi sh from the Twee River, South Africa, where it is threatened by alien fi sh species. © D. IMPSON |

SPECIES IN THE SPOTLIGHT

Jg\Z`\j`ek_\jgfkc`^_k K_\Kn\\I`m\ii\[ÔeÆX:i`k`ZXccp

Skelton, P.H.¹

he Twee River redfi n Heks, Suurvlei and Middeldeur in the Twee River is one of these (Barbus erubescens Skelton) rivers. distinct populations. The Cape was described in 1974, At the time of discovery, only galaxias is currently assessed in the following an investigation one other fi sh species was known IUCN Red List as Data Defi cient, Kthat included extensive fi eld to be indigenous to the Twee River, due to the taxonomic confusion observations. The species is named and both species were isolated by associated with the species for the bright reddish breeding a vertical waterfall of about 10m, complex. Below the falls several dress assumed by spawning located close to the confl uence other indigenous freshwater fi sh males, with females being less of the Twee and Leeu rivers. This species are found, most of them intensely coloured. The common other indigenous fi sh is a species of endemic to the Olifants system. name indicates that the species’ South African Galaxias, formerly One of these species, Barbus calidus, distribution is restricted to one named as the Cape galaxias is the sister species of B. erubescens tributary system of the Olifants (Galaxias zebratus); however, more (i.e., it is the phylogenetically River in the Cedarberg Mountains recently it has become evident most closely related species to of the Western Cape, South Africa. that a number of populations of G. B. erubescens). Barbus calidus, the This tributary system includes the zebratus might represent distinct Clanwilliam redfi n, itself classifi ed Twee and some of its affl uents, the species. The population of Galaxias as Vulnerable, due to threats

1 South African Institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown 6140, South Africa 85 86 CHAPTER 3 | SPECIES IN THE SPOTLIGHT this decline areseveral, including its formerrange. Thereasonsfor and isabsentfrom largesectionsof population hasdeclinedmarkedly zones. Sincethe1970s, juveniles shoalingalongmarginal habitats inpoolsandruns, adults occupyingopenwater the tributarysystem–withlarger was commonandwidespreadin When fi :fej\imXk`feZfeZ\iej other benthicsurfaces. invertebrates orfromrocksand feeds ondriftinginsectsandother up tofi form ofparentalcare.Itcanlivefor water) anddoesnotpracticeany (releasing thegametesinto species isa‘broadcastspawner’ swollen byfrontalrains.The to December)whenstreamsare in springorearlysummer(October several pursuantmales.Thisoccurs over cobblesorrockcreviceswith are attractedandenticedtospawn which individualbreedingfemales school againstrocksurfacesto forming adense,swarming,nuptial features malescongregatingand vestigial serrations. fi and anunbranchedrayinthedorsal of welldevelopedmouthbarbels, tubercles onbothsexes,twopairs such assmallscatterednuptial several otherdistinctivefeatures, other Africanbarbinespecies.Ithas usually, seven–morethanany fi the highnumberofbranchedanal distinctive externalcharacteris chromosomes intotal.Itsmost of eachchromosome),with100 tetraploid (thatis,ithasfoursets Africa, the the temperatereachesofsouthern (80%) ofbarbineminnowsfrom disproportionately largenumber description. Incommonwitha Twee Riverredfi is discussedbelow. degradation causedbyagriculture, from invasivespecies,andhabitat n thatshowseitherincipientor n branchedrays–sixor, more The species’breedingbehaviour Much hasbeenlearntaboutthe ve orsixyears.Thespecies rst discovered,thespecies

Twee Riverredﬁ n sinceitsoriginal n

is the Capekurper( was aSouthAfricananabantid, alien fi sunfi that theTwee River redfi a predatorandgrowsmuchlarger precise impactisnotknown,it is of theTwee and,althoughits mainly inthedownstreamreaches Clanwilliam yellowfi from otherintroducedspecies!The that speciesinthefaceofthreats authorities seekingtoconserve waterfall byNature Conservation the Twee Riverabovethebarrier River system,wasintroducedto a largecyprinidoftheOlifants yellowfi invertebrates. TheClanwilliam predator onsmallfi of thetributaryandanavid is widespreadthroughoutmost which, althoughnotalargefi invasive fi quality andquantity, andalien orchards) impactingbothwater developments (riparianfruit likely impactsfromagricultural also invaded thesystem. Rainbow small fi species, andanother predatoron a North Americancentrarchid The Twee RiverintheCedarburgMountains,Western Cape,SouthAfrica. © SAIAB/P. SKELTON sh ( sh speciestoberecorded shes and invertebrates,have sh ( Lepomis macrochirus sh species.Thefi Labeobarbus capensis Sandelia capensis shes and sh isfound n. Bluegill rst ), sh, ), ) but arenotcommon. been recordedfromtheTwee River be secured. species inSouth Africamight of theseandother indigenous elements areinplace,thesurvival local landowningpublic.Ifthose informed public,especiallythe species forfi may supportintroductionsof by thepublic(who,forexample, in thefaceofcontraryperceptions authorities todowhattheymust action –politicalwillbythe are essentialforconservation might besealedforever. Two things the threatenedindigenousspecies alien speciesistaken,thefateof determined actiontoremovethe sanctuary reachesand,unless alien species.Therearefewnatural pressures aswelltheinvading subject toincreasingagricultural system isrestrictedinsizeand great concern,asthetributary decline intheirpopulationsisof and the conservation statusoftheredfi occasions todeterminethe extensively surveyedonseveral trout ( The Twee Riverhasbeen Oncorhynchus mykiss Galaxias shing), andaproperly species.The ) have n

CHAPTER 3 Jg\Z`\j`ek_\jgfkc`^_k :Xlc[ifej]fiÔj_Y`f[`m\ij`kp1 |

n\jk\ie8]i`ZXËjZiXk\icXb\j SPECIES IN THE SPOTLIGHT McGregor Reid, G.¹ and Gibson, C.¹

lobally, crater lakes are comparatively rare, usually small and specialised freshwater >habitats formed in geological depressions, such as the Ojos del Salado in the Andes mountains,

bordering Argentina and Chile – probably the highest altitude permanent lake of any description (68º32ĿW, 27º07’S, elevation 6,390m, diameter 100m, depth perhaps 5 to 10m). Crater lakes are well represented in tropical Africa, especially in the Guinean rainforest zone of Cameroon, where there Stomatepia mongo, a Critically Endangered cichlid endemic to Lake Brombi Mbo, may be 36 or more. The entire Cameroon. © OLIVER LUCANUS/BELOWWATER.COM region is a celebrated ‘biodiversity hotspot’ for both lacustrine and have attracted much international by vulcanism (Decker and Decker riverine fi shes (Reid 1989; 1996; scientifi c attention. Crater lake 1997; Sigurösson 1999). They are Teugels et al. 1992; Schliewen 2005; cichlids, their taxonomy, phylogeny often visible in photographic, radar Stiassny et al. 2007). Contemporary and ecology were documented early and other imagery taken from space general studies on the world’s on in Cameroon, notably in Lake (Hamilton 2001). crater lakes address important Barombi Mbo (Trewavas 1962; topics such as: lake formation; Trewavas et al. 1972; see below); and Impact crater lakes physical, chemical, geological, they continue to be discovered – for The impact of a meteorite, asteroid geographical and biological example, the recently documented or comet creates a depression. This evolution; paleoecology; historical ‘fl ock’ of eight new species of Tilapia can be a simple bowl (depth to biotic colonisation; and recent from Lake Bermin or Beme (5°9’N, diameter ratio typically 1:5 to 1:7) ecology – including the assessment 9°38’E; diameter around 700m, depth or a larger, shallower, more complex of conservation status and threats around 16m, and age probably far depression (depth to diameter to the survival of the contained less than 1 million years) (Stiassny ratio 1:10 to 1:20) sometimes habitats and species. The potential et al. 2002; Schliewen 2005). Such incorporating a central island or for (and impacts from) human use Cameroonian assemblages are often islands. Such islands are caused is studied, including water supply, regarded as small-scale tilapiine by a gravitational collapse of the agriculture, fi sheries and also counterparts to the better known rim and a rebound of material recreation and ecotourism – such large haplochromine and other to the centre, analogous to the lakes often being scenic locations. cichlid ‘species fl ocks’ of the East splash effect seen when raindrops Crater lakes everywhere may African Great Lakes (Klett and hit water. An island may itself contain a substantial number Meyer 2002; Salzburger and Meyer incorporate a hollow that later of endemic fi shes and other 2004). forms a ’lake within a lake’, as in aquatic and amphibious taxa. Lake Taal, Philippines (Reid pers. Among African fi shes endemic to =fidXk`fe obs.). In geological terms, impact craters, small phyletic and trophic Whatever the location, all craters depressions occur frequently but assemblages of species and genera on earth are formed either by are often temporary, and only representing the family Cichlidae impact of extraterrestrial bodies or some 120 are currently known

1 North of England Zoological Society, Caughall Road, Upton, Chester CH2 1LH, UK 87 88 CHAPTER 3 | SPECIES IN THE SPOTLIGHT ancient, mucherodedimpact western Africa example ofalarge, of northernChad, containsarare and ecologicalcolonisation. erosion, depositionofsediments any underlyingvolcanicactivity, and biologicalprocessesincluding chemical, geological,geographical the depression,andsubsequent the location,scaleandformof nature andpersistenceinclude Other factorsdetermining some laterstageoflakeevolution. become bioticallysignifi silica richglasses.Allofthismay and metamorphism, ‘rockmelt’ such asmeteoriteshards,shock physical andchemicaloutputs, and identifying‘signature’–the of theimpactor, its composition substrate,thevelocity the ‘target’ of impactdepressionsdependon nature, persistenceandeffects (around 60millionyearsago).The since atleasttheCretaceousperiod the marineenvironmentoccurring extinction eventsonlandandin associated withperiodicspecies and biologicaltermsarelikely of importanceinbothgeological impacts, particularlylargeones,are postulated thatmultipleterrestrial considerable scientifi occurrence inAfricaisthereforeof and Australia.Theiroccasional from North America, Europe worldwide, mostcommonly tobetheoldestradiocarbon-datedcraterlakeinAfrica. Lake BarombiMbo,Cameroon.Thislakeisconsidered Aorounga, intheSaharaDesert c interest.Itis cant at aquatic and terrestrialpatterns in determining much ofthelarge-scale Mega Chadhas beencrucialin human abstractiondemands.The the 1960s,mainlyduetoexcessive is nowonly5%ofitsvolumein few thousandyearsago.LakeChad persisting tosomeextentuntil a years agoto10,000ago)and Holocene periods(around2million dating fromthePleistoceneto more extensive‘MegaLakeChad’ of themorepermanentandfar series, whichmayhavebeenpart The Aroungacraterisoneofalocal of kilometres(Todd carried forhundredsorthousands 1,200,000 tonnesofdustperday in winter, amountto anaverageof the BodéléDepressionandwhich, to duststormsgeneratedwithin fi are usuallydryandcontributea pers. obs.).However, thecraters appear ‘as iffromnowhere’(Reid (frog andtoad)tadpoleswhich inundations ofwater;andanuran resting inthesandsinceprevious shrimps’) emergingfromeggs anacostracan crustaceans(‘fairy of life,albeitbriefl contain aremarkabledensity Sahelian regionsuchpoolsmay pools inrainyperiods.Acrossthe supports isolatedtemporary years ago(Hamilton2001))which 17km; agearound200million crater (19°6’N, 19°15’E;diameter ne diatomaceouslakesubstrate y, including et al . 2007). Federal RepublicofGermany2011). (Hamilton 2001;Embassyofthe nearby intheGulfofGuinea Coast andtooceanicdeposits, Bosumtwi havespreadtotheIvory sediments associatedwithLake Shield andresearchindicatesthat bedrock oftheWest African Africa, itliesovercrystalline natural lakeinsub-Saharanwestern million years).Thelargestsingle maximum depth75m;age1.3±0.2 01°25’W; rimdiameter10.5km; impact craterlake(06°32’N, a comparativelyyoung,permanent known, butstillscarce,exampleof (Reid 1996). Sudan ichthyologicalprovince for westernAfricaandtheNilo- historical andrecentbiogeography MfcZXe`ZZiXk\icXb\j contact withgroundwater. in asubterranean chambermakes when hotvolcanic lavaormagma created inagaseousexplosion 1997). Therockyrimisoften complex (DeckerandDecker of thevolcanicactivitycanbe not alwaysobvious,andthenature However, thesedistinctionsare are shallowerwithalowprofi inverted cones; classes: sometimes dividedintotwo and theirassociatedlakes,are Craters formedthroughvulcanism, Lake Bosumtwi,Ghanaisabetter calderas

whicharedeep and maars ©U.SCHLIEWEN .

which le.

CHAPTER 3 |

SPECIES IN THE SPOTLIGHT

By contrast, Lake Barombi only three known contemporary ‘exploding’ and periodically lethal Mbo is small (see above) and lakes, all of which are African estimated to be biologically (the others being nearby Lake Monoun, Cameroon (5°35’N, mature since about 25,000 to 10°35’E) and Lake Kivu, Rwanda). Nyos and Monoun are located 33,000 years ago; it is considered within the Oku Volcanic Field to be the oldest radiocarbon- near the northern boundary of the Cameroon Volcanic Line, a zone dated crater lake in Africa of volcanoes, maars, calderas and other tectonic activity that extends Subsidence of materials creates a permanently fi ll up with water south-west to the large, inactive depression within the rim that may from snow, rainfall, groundwater, Mount Cameroon composite later fi ll with water. A diatreme a captured drainage, spring or volcano (stratovolcano) and beyond often persists under the lake bed, swamp or a larger inundation. to the island of Bioko in the Gulf that is, a pipe-like vertical volcanic Depending on water supply, of Guinea, which also contains an vent that is fi lled with broken and drainage and evaporation, the unexplored crater lake (Flesness, cemented rock created by a single lake may reach the lowest point pers. comm.). Nyos has periodically explosion. Such diatremes may on the rim and then overspill as a been supersaturated with carbon remain active. Lake Nyos (around waterfall if the rim is high; or as a dioxide (CO2, forming carbonic 322km north-west of Yaoundé, stream, if at the outset the rim is acid) leaching from the underlying Cameroon, close to the border with low or becomes water eroded. At a magma and with a peak lake Nigeria) is an example of a simple critical point of attrition there can density of approximately 90 million maar lake, but a comparatively be catastrophic breakout fl ooding. tonnes of CO2. In 1986, there deep one (6°26Ŀ17ŀN, 010°17Ŀ56ŀE; If the crater contains an active was a gaseous explosion, perhaps 1,091m above sea level; 2km long by volcanic vent (see ’diatreme‘ above) precipitated by an earthquake or 1.2km wide; and 208m maximum the water will have an elevated landslide, releasing approximately depth). Lake Barombi Mbo in temperature and be turbid and 1.6 million tonnes of CO2 into south-west Cameroon is formed in acidic from high concentrations the atmosphere. This killed some a caldera, albeit a fairly small one of dissolved volcanic gases and 1,800 people, 3,500 livestock, and (4°39’46’’N, 9°23’52’’E; 303m above distinctly green, or red-brown if gas in solution presumably killed sea level; 2.15km wide; and around iron rich. Gases include carbon fi shes and other aquatic life.

110m maximum depth) (Schliewen dioxide (CO2), sulfur dioxide (SO2), Degassing pipes were installed 2005; Lebamba et al. 2010). hydrogen chloride (HCl) and in 2001 to prevent a repetition of hydrogen fl uoride (HF), which may the catastrophe (Kling et al. 2005). CXb\[\m\cfgd\ek persist in solution and are lethal to Some 2,000 times larger than Nyos, Whether formed by impact or invertebrate and vertebrate life. Lake Kivu has also been found to vulcanism, craters that persist Lake Nyos, with a diatreme some be periodically supersaturated – anywhere may periodically or 80km below the lake bed, is one of with evidence for outgassing every 89 90 CHAPTER 3 | SPECIES IN THE SPOTLIGHT craters, for example,the lakethat of watercanform veryquicklyin that substantial permanentbodies there iscontemporaryevidence (Schliewen 2005).Inanyevent, as around1millionyearsago of thelakehasbeenestimated et al Africa (Elenga radiocarbon-dated craterlakein ago; itisconsideredtobetheoldest since about25,000to33,000years estimated tobebiologicallymature Lake BarombiMboissmalland of over1,000km².Bycontrast, 70,000 yearsago,withanarea Indonesia wasformedaround the LakeToba caldera,DanauToba, maturity andscale,forexample, crater lakesareofconsiderable other limnologicalprocesses.Some overturns ofthermalstrata),andby (including infl 2010), nutrients,watermovements (Elenga and biotaabovethewaterline the natureofcraterrimsoil high, butthisisdeterminedby and primaryproduction)canbe level ofphotosyntheticactivity hence transmissionofsunlight, The clarityortransparency(and can, insomeinstances,besaline. life supportingandfreshbut down toaround40mareusually biomass. Lakesurfacewaters the animalandplantspecies layers whichsupportmostof warm, lessdense,sunlitsurface is usuallyquiteseparatefromthe anoxic waterabovethelakebed The deep,cold,dense,aphoticand eventually ecologicallystratifi normally becomesthermallyand dormant volcanocraters,thewater impact cratersandinactiveor other aquaticanimalpopulations. ecology andevolutionoffi and indeterminingthesurvival, sequestration and‘footprints’– stability, contemporarycarbon important whencalculatinglake periods andalsoreleaseitisclearly carbon dioxideatdepthforlong ability ofcraterlakestostore thousand yearsorso.Thegeneral In thecaseoflargemature . 2010).Thephysicalorigin et al . 2004;Lebamba ows, outfl et al . 2004;Lebamba ows and shes and et al ed. . their endemiccichlidfi conservationists mainlybecauseof of evolutionarybiologistsand lakes haveattractedtheattention African andothersmallcrater speciation oradaptiveradiation are African models oftilapiinecichlid possibilities aside, western test scientifi and diffi humans orbirdsarelesscredible introductions offi extant). Notions ofpaleo-historical crater streamoutfl indundations ofwater, orvia through large-scalepaleo-historical hypotheses ofnaturalmigration there arepotentiallytestable crater isintriguing.Insomecases, of howtheycametooccupythe geologically isolated,thequestion contain cichlids,andwhichare (Schliewen 2005).For thosewhich contain fi all westernAfricancraterlakes occupied byinvertebrates,not and speciation.While invariably of populationdifferentiation environment, andtheprocesses from aninitiallylifeless stages inecologicalcolonisation an opportunitytoinvestigate Great Lakes.Suchcratersprovide easily studiedthantheEastAfrican ecosystem –a‘microcosm’more (if potentiallymorevolatile) represent ayounger, lesscomplex to theirsurvival.Thecraters natural andanthropogenicthreats evolution ofspecies Lake colonisationandthe Philippines. the eruptionofMountPinatubo, developed post1991,following African Great Lakes easily studiedthantheEast ecosystem –a‘microcosm’ more potentially more volatile) cult, orimpossible,to shes andshrimps The cratersrepresent a younger, lesscomplex(if cally. Settingsuch shes oreggsby ows (somestill . Western shes andthe and Meyer2004;Seehausen2006). (Klett andMeyer2002;Salzburger grand-scale easternAfricanmodel being testedagainsttheclassical these competing (butnot sympatric speciation. Intesting that is,through aprocessof without suchtotalisolation, fl species canariseasmonophyletic with theircongeners. ,but bythattimedidnotinterbreed high paleo-historicalwaterlevels returned tothemainlakeduring These isolatessupposedlylater genetically intodistinctspecies. morphologically, behaviourallyand then differentiatedecologically, isolate cichlidpopulations,which time createdsatellitelakesto and fallofwatersingeological was suggestedthataregularrise separation ofpopulations.It involving periodicgeographical allopatric speciation,thatis,one hundreds throughaprocessof species fl cichlid taxaevolvedintolacustrine is thathaplochromineandother general easternAfricanhypothesis initially onLakeVictoria, the water ordeepwaterhabitats.’Based species-rich andliveintheopen fl the oldestlineagesofaspecies age, ofthelakes.We observethat with thesurfacearea,butnot seems toberoughlycorrelated Meyer (2004):‘Speciesrichness However, forSalzburgerand or confi enclosed, lacustrinephenomenon species fl ocks withinthebodyofa lake ock ofcichlidsareoftenless Evidently, theevolutionof An alternativemodelisthat ned tocichlidtaxa. ocks numberinginthe ocks isnotinvariablyan CHAPTER 3 necessarily mutually exclusive) standard recognised freshwater deforestation and slash and burn models, Schliewen et al. (2001) ecoregions of the world and Africa agriculture within the crater rim); conducted a ‘gene fl ow’ study (Thieme et al. 2005; Abell et al. adverse water level fl uctuation within fi ve tilapiine morphs 2008). (from damming the lake outfl ow endemic to Lake Ejagham, Thieme et al. (2005) designate and from excessive abstraction); | western Cameroon (5°44’59”N, closed basins and small lakes and water pollution (from natural SPECIES IN THE SPOTLIGHT 8°59’16”E; surface areas 0.49km2; as a ‘major habitat type’, whose volcanic gases, from aerial and maximum depth around 18m ultimate conservation status within industrial emissions travelling (Schliewen 2005)). Comparisons most of the western African block from a distance; and from locally with a closely related riverine of ecoregions is under threat ‘based applied agrochemicals, pesticides outgroup of cichlids suggest that on projected impacts from climate and ichthyotoxic molluscicides synapotypic colouration and change, planned developments, used to control the aquatic snail ‘differential ecological adaptations and human population growth’. vectors of human schistosomiasis, in combination with assortative Recent research on pollen, biomes, at least endemic in Barombi Mbo). mating could easily lead to forest succession and climate in Among conservation speciation in sympatry’ (Schliewen Lake Barombi Mbo crater during recommendations that have

et al. 2001). More generally, it the last 33,000 years or so suggests been proposed by the authors is postulated that a dynamic the persistence of a humid, dense, (above) are: systematic Population network of gene exchange or evergreen cum semi-deciduous and Habitat Viability Analyses, hybridization among populations forest: ’These forests display a as formulated by the IUCN creates a process of ‘reticulate mature character until ca 2800 cal Conservation Breeding Specialist sympatric speciation’ among yr BP then become of secondary Group; Red List threat assessments Cameroonian crater lake cichlids type during the last millennium (as summarized in this volume); (Schliewen et al. 1994; Schliewen probably linked to increased the formal designation of lakes as 1996, 2005; Schliewen and Klee human interferences [our emphasis]’ legally and practically protected 2004). Comparable empirical (Lebamba et al. 2010). aquatic nature reserves of national research on post-colonisation The recent conservation status and international importance, with cichlids in a young crater lake in of small Cameroonian crater an accompanying conservation Nicaragua also supports the idea lakes, including Barombi Mbo, action plan (Lakes Barombi Mbo that sympatric endemic ‘morphs’ and their endemic fi shes and and Ejagham have now been of individual cichlid species may invertebrates, is considered in designated as forest reserves diversify rapidly (say, within a detail by Reid (1989, 1990a,b, 1995, (Schliewen 2005)); and ex situ hundred years or generations) in 1996) and Schliewen (1996, 2005). programmes for the conservation ecology, morphology and genetics Such unique lake environments breeding of species at risk, and this can be interpreted as and endemic species are clearly with the prospect of eventual ‘incipient speciation’ (Elmer et al. of national and international reintroduction in appropriate 2010). Again, this is postulated to importance. There is, from the circumstances (such ex situ be through disruptive selection, outset, an inherent vulnerability aquarium breeding programmes perhaps sexual selection, mediated of these ecosystems resulting have been in operation since 1999 by female mate choice. from the geological instability in through European and North craters; their small physical size; American Fish Taxon Advisory Conservation of crater lake fi shes. the small size of the contained Groups). Despite the persistent The phylogenetic and associated populations and their genetic threats outlined above, a survey of data on crater lake cichlid species isolation; and, for cichlid fi shes, Lake Barombi Mbo in 2002 found fl ocks (above) are at different their methods of reproduction all fi sh species to still be present levels of generality and, among and limited fecundity. Actual or (Schliewen 2005). However, other criteria, important in potential general threats are widely many of the species present the evaluation of conservation familiar, including: overfi shing are threatened (even Critically priorities (Stiassny and de Pinna and other socio-economic factors, Endangered), but there have been 1994). However, a paucity of including pressure from external no recorded fi sh or invertebrate well-worked and wide-ranging visiting tourists; the introduction population declines to the point studies has until recently limited of alien species (for example, of extinction in any of the crater such contributions (Stiassny crustaceans and fi shes (Slootweg lakes. Nevertheless, continued 2002; Stiassny et al. 2002). Even 1989)); siltation and a reduction or vigilance, conservation monitoring, so, western African crater lakes loss of allochthonous food supply threat assessment, mitigation and are included as an important of terrestrial plant material and protective measures remain highly biogeographic category within invertebrates (both resulting from appropriate. 91 Chapter 4. Freshwater molluscs of Africa: diversity, distribution, and conservation

Seddon, M.¹, Appleton, C.², Van Damme, D.³ and Graf, D.4

1 Bracken Tor, Saxongate, Okehampton, Devon, EX20 1QW 2 School of Biological and Conservation Sciences, Westville Campus, University of KwaZulu-Natal, Durban 4000, South Africa 3 University of Ghent, Sint-Pietersnieuwstraat 25, B 9000 Ghent, Belgium 4 Department of Biological Sciences, University of Alabama, Tuscaloosa AL 35487 USA

The Kulungu River, part of the Chambeshi basin in the Upper Congo. © DANIEL GRAF AND KEVIN CUMMINGS

Diving for mussels in the Upper Chambeshi River, Upper Congo. © DANIEL GRAF AND KEVIN CUMMINGS CONTENTS 4.3.1 Regional patterns of threat 112 Introduction 94 4.3.2 Dams 113 4.1 Overview of freshwater molluscan diversity in Africa 96 4.3.3 Water pollution 114 4.1.1 Molluscan diversity in river basins 97 4.3.4 Sedimentation 114 4.1.2 Molluscan diversity in ancient lakes 100 4.3.5 Invasive species – water hyacinth and other plants 114 4.1.3 Molluscan diversity in artesian basins 102 4.3.6 Invasive species – ﬁ sh farming 114 4.1.4 Molluscan diversity in saline lagoons, saltmarshes and mangroves 102 4.3.7 Invasive species – molluscs and their impact on indigenous species 114 4.1.5 Molluscan diversity in crater lakes 103 4.3.8 Climate change 117 4.1.6 Molluscan diversity in montane lakes 104 4.4 Understanding the impact of molluscs on 4.2 Conservation status and distribution of human health and livelihoods 117 freshwater molluscs 104 4.5 Recommended conservation measures 118 4.2.1 Threatened species 105 4.5.1 Recommended research actions 119 4.2.1.1 Changes since the 1996 assessment 106 Species in the spotlight – Freshwater molluscs are 4.2.2 Restricted range species 106 important – especially to the African openbill stork 120 4.2.3 Data Deﬁ cient species 109 Species in the spotlight – Can the last populations of the 4.2.4 Extinction patterns 110 giant African freshwater pearl mussel be saved? 123 4.2.5 Undescribed species 111 Species in the spotlight – A mollusc with a bit 4.3 Main threats 112 of muscle 124

Children washing clothes in a steam near the Sanaga River in Cameroon, potentially exposed to diseases hosted by freshwater molluscs. © KEVIN SMITH 94 CHAPTER 4 | MOLLUSCS north-eastern Africa, central Africa and western Africahave Africa, centralAfrica andwestern north-eastern studies forMozambique (Germain1935).Incomparison monographs (e.g., Connolly1939)aswell Germain’s throughout thelasttwo centuries,withConnolly’s various regions, Africa, havereceived suchassouthern attention molluscs from regions Africa.Some includingwestern malacologists inAfrica,anddescribed someofthefi continent (Brown 1994). Adansonwasoneoftheearliest during theperiodsofcolonialexpansionthroughout the Exploration ofAfricanfreshwater molluscscommenced accompanying DVD. by taxonomicgrouping ratherthanregion, pleaseseethe conservation actions.Forasynthesisofresults presented highlighting theregions andspeciesthatrequire further an overviewofthethreats tothecontinentalfaunaand for theentire pan-Africanmolluscanfauna,providing 2010; Graf et al. were determinedthrough aseriesofassessments(Darwall andcentralAfrica northern western, southern, eastern, From 2003to2009,theregional statusofmolluscsin over thenextfi have anincreased numberofendemicspeciesdescribed (West Damme is rapidlychangingthenumberofknownspecies(Van survey, combinedwithreview ofhistoricalcollections, some generaalready. Africa,theleveloffi Innorthern species maywellmultiply, ashashappenedinstudiesof utilising molecularsystematics–thenumberofknown that astaxonomicreviews continue–especiallythose of attentionasEuropean andNorthAmericanfaunas, freshwater molluscshavenotyetreceived thesamelevel prosobranchs. We needtorecognise thatthetropical concentrated onthefreshwater Unionidmusselsandthe Most research efforts inrecent yearshave,however, proportion ofthewidespread, more cosmopolitanspecies. Gastropoda, withthepulmonatescontainingahigher the pulmonates.TheBivalviaare lessnumerous thanthe dividing intotwoinformalgroups, theprosobranchs and groups, theBivalviaandGastropoda, withthelatter Bogan 2008).Freshwater molluscsfallintotwo main are about780European species)(Seddonpers.comm.; around 1,540speciesforthePalaearctic (ofwhichthere compared toaround 880speciesforNorthAmericaand America andEurope, withonlyanestimated560species diverse thansomecontinentalfaunassuchasinNorth In general,thefreshwater molluscsofAfricaare less for understandingourchangingfreshwater diversity. interactions, allofwhichunderscore theiruseasproxies life-history strategiesandexhibitcomplexecological found inawiderangeoffreshwater habitats,havevaried Freshwater molluscs(bivalvesandgastropods) are Introduction 2005;Kristensen et al. et al. et al. 2003),andtheseareas canbeexpectedto 2010),asisalsothecaseinLakeTanganyika ve years. 2011).Thischaptersynthesisestheresults et al. 2009a,b;Van Damme et al. eld rst

for their overallpan-African status.Most ofthesewould regional assessments,andhencecould notbeevaluated project considered valid,that were omittedinthe earlier As such,there are somespecieswhichthepan-African with thatof concept ofthegenus Regardless, whilethe namehaschanged,thetaxonomic relating thatmaterial to livingforms(Graf,pers.comm.). shells, andneontologiststookaconservativeapproach to Chambardia, confusion seemstohavebeenthatthetypespeciesof priority (Vaught 1989;Daget1998).Thebasisforthe recognized thatthename species circumscribed underthistaxon,ithassincebeen referred tothegenusas Through thelate1990s,Mandahl-Barth(1988)andothers assigned indifferent waysinthevariousregional reports. in freshwater mussels,thegenus as theinitialbasisfortaxonomiclist.Forexample, this project havediffered depending onwhichwastaken (Graf pers.comm.).Assuch,theregional treatments in on opinionsofthevariabilityformspeciesshells lumping, butneitherisbasedondefi the nature ofthetwotaxonomicschoolssplittingand rather different viewsofthestatusbivalves,refl and Mandahl-Barth(1988).Thesevolumesprovide two most activitypresented inthecataloguesofDaget(1998) By contrast,there are fewstudiesoftheBivalvia,with focus ofvariousstudiesoverthelast30years. are pulmonates,andassuch,thesetaxahavebeenthe papers andsmallhandbooks.Manyofthehostspecies young scientistsinAfrica,andproduced manyresearch many trainingprogrammes supportingagenerationof established theDanishBilharziaLaboratory, whichhosted of thefreshwater gastropods todate.Mandahl-Barth remains themostcompletehandbooktoidentifi Barth andD.Brown. Brown’s monograph(Brown 1994) the majorityofworkinearlydays,Mandahl- medical importance.Two notable scientistscontributed attention fortaxonomicstudiesrelated tomolluscs of Africa has,forthelast50years,alsobeenfocusof efforts are lookingattheoriginsofotherpulmonategenera. subfamily intheFamilyPlanorbidae),andfurtherresearch is revealing more ontheoriginsofAncylinae(nowa prosobranch families.Recentresearch byC.Albrecht incorporating molecularsystematicscontinuesonselected been produced (West revised handbooktoGastropods ofLakeTanganyika has volumes onEastAfricanlakefaunas.More recently, a E. Leloup(1935)andG.Mandahl-Barthbothpublished as theresult ofmore recent surveywork.Inthe1950s, Smith andK.vonMartenshavebeenconsiderablyrevised studies ofLakeTanganyika by researchers suchasE.A. to theunusualfaunaofancientRiftValley. Theearly Africanfaunareceivedeastern more attention,partly due received relatively littleattentioninthelast50years.The

Spathopsis C. letourneuxi et al. , soconfusionshouldbelimited. Chambardia Spathopsis Chambardia , wasdescribedfrom (sub)fossil 2003), andfurtherresearch Chambardia . However, giventhe isinterchangeable nitive data,mainly hasnomenclatural hasbeen ecting cation CHAPTER 4 | MOLLUSCS 95

ed snail. Coelatura , ve species not Mutela legumen 2001; Cummings (LC), et al. M. “rostrata” (LC), nal pan-African assessments. (LC) and an unidentifi shes (Wächtler Chelidonopsis hirundo C. gabonensis (VU), Aspatharia pfeifferiana (VU), rotula Unionid mussels collected from Malebo Pool in the Republic of the Congo; © OLIVERLUCANUS/BELOWWATER.COM evaluated) occupy a wide range of habitats, from brackish from evaluated) occupy a wide range of habitats, environments, estuarine waters to interior lentic and lotic cosmopolitan taxa in the pea- more are although there bivalves possess clams (Sphaeriidae). All of the freshwater These a common suite of adaptations to life in freshwater. development, and, in the direct include larval brooding, mussels, obligate larval parasitism case of freshwater fi upon freshwater derived and Graf 2009). Despite these common characters and Ancylinae. There are also a number of introduced of introduced a number also are There and Ancylinae. evaluated not been that have Physidae in the family species native range. Africa is outside their given that bivalves Freshwater 25% of approximately bivalves represent The freshwater continent. In mollusc fauna of the African the freshwater nine families, 158 species (28 genera, total, we recognise 17 species that of bivalves in Africa, including two orders) due to differences not evaluated as part of this project were assessments that in the regional in taxonomic treatments the basis for the fi provide distributed orders: divided into two globally The bivalves are mussels) and known as freshwater Unionoida (otherwise pea-clams). (otherwise known as clams and Veneroida mussels contain the higher number of species Freshwater strictly freshwater, (82 species). All unionoid species are common and often locally and in Africa these molluscs are lakes. Similarly, abundant inhabitants of both rivers and including fi (56 species, the Veneroida cient or Least Concern. © HEINZ H. BÜSCHER (LC), a species endemic to Lake Tanganyika, (LC), a species endemic to Lake Tanganyika, be assessed as Data Defi family to family and This variability in knowledge from a considerable challenge presented between regions has research to this assessment and, although ongoing our knowledge of the fauna, there considerably increased need for continued investment and a pressing remains knowledge base. training to build on the current gastropods Freshwater about 75% of all represent gastropods Freshwater molluscs in the African continent, dividing freshwater and the the prosobranchs into two informal groups, the higher (2005) revised pulmonates. Bouchet and Rocroi could phylogeny of Mollusca, noting that “Prosobranchia” no longer be supported as a formal designation, as it was polyphyletic, with multiple origins within the group. two major clades, the Caenogastropoda are Now there along with smaller clades such and the Heterobranchia, “Pulmonata”, as Neritimorpha. Within the informal group have also been found, however most of similar problems lie in the Basommatophora, within species the freshwater (1994) estimated that there the Eupulmonata. Brown species have been in Africa, but more 326 species were study as new species have been current in this recognised described and molecular systematics allowed some taxa to be revised. lakes most varied within the are prosobranchs Freshwater pulmonates freshwater river systems, whereas and larger and often associated with small water-bodies more are of seasonal drying. In total, 353 species tolerant many are and 143 species of pulmonates are of prosobranch Planorbidae, in the families Lymnaeidae, here, recognised where impacted by sedimentation, dredging it is locally and collection for the shell trade. Tiphobia horei Tiphobia horei 96 CHAPTER 4 | MOLLUSCS patterns of dispersalandabundance. patterns of theirdisparatelifehistories,demonstratearange represent different evolutionary lineagesand,asaresult from shared environmental pressures, thesebivalvetaxa provide ageneralised mapof theprojected hotspots of and Gargimony (in Groombridge and Jenkins 1998) can befoundin thedifferent regions ofAfrica. Bouchet artesian basins(Seddon2000), andallofthesehabitats and diversityinancientlakes, large riverbasinsand Freshwater molluscs fi 4.1 compiled byBouchetandGargimony, adaptedfrom Groombridge andJenkins1998). Figure 4.1.Hotspotsoffreshwater molluscdiversityacross continentalAfrica(highlightedinorange)(data diversity inAfrica Overview offreshwater molluscan nd theirhighestlevelsofendemism shared withthe Palaearctic fauna, the Arabianfauna and Africawhichhaselements The exceptionis north-eastern south oftheSahara where anAfrotropical faunadominates. the faunalieswithPalaearctic fauna,andtheregions into theregions north oftheSahara,where theaffi African freshwater mollusc faunaisgeographicallydivided species distributions,ratherthan experts’bestestimates. this picture toberefi species diversityofAfricaasdefi an in-depthcomparison(Figure 4.2)ofthehotspots The outputsofthecurrent pan-Africanproject nowallow Lakes and the river rapids of western Africa Lakes andtheriverrapidsofwestern various partsofAfrica,namelytheEastAfricanRiftValley molluscan freshwater diversityintheworld,highlighting ned onthebasisofactualmapped ned in1998,andenable (Figure 4.1) nity of nity . CHAPTER 4 | MOLLUSCS 97 Burnupia and 2005). Brown (1994) (1994) 2005). Brown Ferrissia et al. uvial mollusc faunas of these cient genera 4.1.1 Molluscan diversity in river basins the Nile systems in Africa are freshwater The two largest the Congo and the Congo rivers. The habitats of the largest, less diverse, consisting mainly of rainforest the are River, rift lakes (Thieme rivers and large the Bocxlaer (2009) compared Damme and Van and Van species richness of the fl lakes. Brown two river basins, with exclusion of the large the known from pointed out that the number of gastropods Nile Basin (51) is about half that of the Congo (96) (if we take the taxonomically defi of basin endemics is as genera only), and the proportion for the namely 10 out of 51 species. Similarly, much lower, bivalves, species diversity is lower in the Nile Basin, with a Lower Congo . 2010). The southern et al. Malebo Pool in the Congo Basin Additional regions of interest not previously not previously of interest Additional regions The regions highlighted in this study as having the highest highlighted The regions 4.2) include the East African species diversity (Figure and Lake Malawi), Lake Tanganyika Lakes (Lake Victoria, Lake Mweru, the the Upper Congo catchments around of western river rapid regions Africa, and the highlighted in Bouchet and Gargimony’s analysis (Figure (Figure analysis highlighted in Bouchet and Gargimony’s 4.1) include the North African spring-snails, the East the Lake Chad African coastal rivers, Lower Nile River, basin and Rapids. the African fauna (Van Damme the African fauna (Van fauna is generally drawn through line for the Palaearctic Libya and the Ahagger Mountains, north-east towards 1961). west to Mauritania (Sparks and Grove Figure 4.2. Patterns per river/ of species number richness = Species species. all mollusc for richness of species Figure lake sub-catchment. 98 CHAPTER 4 | MOLLUSCS The KulunguRiver, partoftheChambeshibasininUpperCongo. some rare andrangerestricted species. Forexample Congo Basin 5.3 millionyearsago). ofAfricainMiddleandLateMiocene times(23to Horn main invasionoffreshwater molluscsintoAfricaviathe Damme limited numberofendemicswith aPalaearctic origin.Van limitsoftheirrange,alongwitha species atthesouthern of thePalaearctic faunaoccur, includingwidespread zone (Morocco, Algeria,Tunisia andLibya),components Nile andintheEthiopianHighlands,aswithMaghreb Corbiculidae) are more abundant.IntheLower (Egyptian) clams andpeamussels(FamiliesSphaeriidae while intheNileBasinpulmonatesnailsandsmaller and unionoidbivalvesare overwhelminglydominant, temporary pools.IntheCongoBasinprosobranch snails Basin isofgeneralisttaxalivinginfringehabitats,suchas in theriversandlakes,highestdiversityNile richness oftheCongoBasinisforspecialisttaxathatlive where thisdiversityisfound.Whilethehighestspecies Bocxlaer (2009)alsoobserveddissimilarityinthehabitats 1998; GrafandCummings2007a,b).Van DammeandVan Nile Basinascompared to18intheCongoBasin(Daget in theNileBasinwithonlyfi and thedegree ofbivalveendemismisdramaticallylower total of28speciescompared to38intheCongodrainage, et al . (2010) suggest this pattern stemsfrom. (2010)suggestthispattern the : Thefaunafrom theCongoBasinincludes ve speciesendemic tothe speciose, butonlyone species, family Thiaridae theAfro-Asian genus Africa(Vanto eastern DammeandPickford 2003).Inthe distribution andthatthegenus’ rangeformerlyextended these basins.Fossilevidence shows thatthisisarelict and theVolta River, anditsrepresentatives are rare within genus and identifi as CriticallyEndangered duetotheirrestricted ranges River. Allrepresentatives ofthesethree generaare listed are restricted tothefastfl Pseudogibbula ranges from SomaliatoMozambique(fourspecies),whilst as itisspread bybirds), another( by Brown (1994)is Assimineidae, oneofthefi species listedasCriticallyEndangered. Inthefamily shallow fastfl habitats, where itliveslikealimpetfi the KouilouRiver(RepublicofCongo),inhabitssimilar neritoides, Africa. InthefamilyAmpullaridae,anotherspecies, are probably relicts, asare thesimilartaxainwestern exclusively foundintheLowerCongoarea. Thesegenera adapted tolifeonstonesinfastfl Gongodoma, Liminitesta there are three monotypicgenerainthefamilyBithyniidae, © DANIELGRAFANDKEVINCUMMINGS Pseudocleopatra whichwasrecently discovered inatributaryof ed threats. InthefamilyPaludominidae, owing water. Thisspeciesistheonly , Valvatorbis Assiminea isrestricted totheLowerCongo and owing partoftheLowerCongo ve generarecognized inAfrica and (whichisfairlycosmopolitan, Septariellina Funduella, Melanoides tuberculatus owing streams, andare Eussoia) Melanoides xed onrocks in whichare all isAfricanand (fourspecies) ishighly Lanistes Lanistes ,

CHAPTER 4 | MOLLUSCS 99

c (Mandahl- is limited to Nilo- is limited Coelatura © JEAN-PIERRE BOUDOT Nitia cient. The Oued Za in Morocco, a typical river of the Maghreb. owing, highly oxygenated are listed from Africa (plus listed from are Unio, Anodonta, Leguminaia Nitia The freshwater mussels of this region mussels of this region The freshwater is a widespread Eurasian and North is a widespread that inhabit the Palaearctic region of region that inhabit the Palaearctic Anodonta Potomida as a morphologically distinct group of species, worthy of distinct group as a morphologically conservation actions to safeguard the few rare and unique the few rare conservation actions to safeguard species occurring in the fast-fl waters of the river rapids. North-west Africa: within the genera of largely are The Niger River basin contains fewer endemic species The Niger River basin and Nile river molluscs than the Congo of freshwater the Senegal, other rivers such as systems. Similarly, species whilst having reasonable Orange, and Zambezi, species. Some of few endemic have relatively diversity, specifi require these western do, however, African rivers and The of Europe. northern and parts Africa, the Maghreb, genus reaches American genus that, in the western Palaearctic, one that was not evaluated), all of which are assessed as all of which are one that was not evaluated), either Least Concern or Data Defi Sudanic Africa, including the Senegal, Niger, Chad, and Chad, Niger, including the Senegal, Sudanic Africa, been have frequently The species of this genus Nile Basins. under the catchall genus, subsumed 2004), but we regard Scholz and Glaubrecht Barth 1988; Nitia and Cummings (Daget 1998; Graf of separate recognition species of 2007a). Three The highest species richness is found in Cameroon, with with in Cameroon, is found richness species The highest Within the threatened. are most of which species, seven mussels the genus freshwater

2008). Sierraia Potodoma et al. (which was moved from moved from (which was owing rivers draining the west owing rivers draining the west ection of the area’s high diversity. high diversity. ection of the area’s Potadoma The western African region is recognised is recognised The western African region owing streams in Sierra Leone and Guinea, owing streams (family Bithyniidae), that are adapted to life on (family Bithyniidae), that are 30 parthenogenetic species are restricted to the to the restricted are species 30 parthenogenetic ca 2005). In this region and in the Congo, the 2005). In this region 1998), and the presence of some unique Prosobranch Prosobranch of some unique 1998), and the presence Soapitia owing rivers that drain the west African plateaus (Thieme owing rivers that drain the west African Africa plateau is a refl Thiaridae to Pachychilidae on the basis of molecular the basis of molecular to Pachychilidae on Thiaridae the found in 2002)) are and Glaubrecht (Köhler research and also in easternrapids of the Lower Congo and north- threatened. eastern Congo, and some of these are Africa: Western biodiversity (Mittermeir of terrestrial as one of the hotspots et al. fast-fl genera in the short, west Africa plateau have localized The rivers draining the genera, species, including two range restricted large African lakes and the Congo, with the highest species with the highest species lakes and the Congo, African large 12 species are where in the Congo Basin, richness found Species of recorded. is found throughout Africa (and Asia), while most of the of the while most (and Asia), Africa throughout is found other and stones in fast-fl and these genera are considered to be threatened as well to be threatened considered and these genera are in origin. The as being potentially quite ancient and relict fast- smaller, River Niger is less diverse than many of the fl et al. species in the family Pachychilidae has a disjunct dispersion species in the family Pachychilidae has a disjunct Congo and Liberia to Lower (in the Atlantic rivers from also in eastern and north-eastern Congo). This ancient as reaching taxon formerly had a much wider distribution, Bocxlaer Basin (Van far east as the Turkana 100 CHAPTER 4 | MOLLUSCS species, from thecontinent,three ofwhichare threatened. Fournew Africa.Intotal,four southern and eastern, but withawiderdistributioninnorthern, Endangered. rivers theyinhabit,bothspeciesare rankedasCritically their restricted distributionranges,andthethreats tothe represent endemicspecies Africanpopulationsmay suggests thatthesenorth-western work (Van Damme methods andspeciesconcepts(Graf2007).More recent only justbeguntobere-evaluated analytical usingmodern are holdoversfrom earlyinthelastcentury, andtheyhave the traditionalconceptsofPalaearctic generaandspecies into RussiaandCentralAsia(Araujo Palaearctic, extendingeastfrom theBritishIslesandIberia morphologically variableandwidespread inthewestern species, conspecifi has beenthatthepopulationsinMorocco andAlgeriaare species of is disagreement amongmalacologistsastowhichtwo Maghreb are represented bytwospecies,althoughthere Africa.Populationsinthe limitinnorth-western its southern Table 4.1.Freshwater gastropod faunaofthelargestAfricanlakes(Based onBrown 1994andadditionaldata): andcentralAfrica valleys thatliebetweeneastern In Africa,theancientlakesare mainlylocatedintherift 4.1.2 Molluscandiversityinancientlakes as welltheTurkana Basin. Unio latter speciesisthemostwidespread ofthePalaearctic- African endemicspecies, regard theAfricanpopulationsof U. foucauldianus populations of other species.Here wetreat African thenorth-western project, beingonlyrecently removed from synonymywith U. ravoisieri the adjacentIberianpeninsula(Araujo Maghreb Africa;allfourare ofnorth-western alsofoundin mancus aeTraa3572041subspecies 1subspecies 0species 4subspecies 4 7 8species/subspecies 8species/subspecies 10 3species/subspecies 15 15 17species 16 7,200 12species/subspecies 21 2,700 65species 27 3,156 2,150 28 4,850 5,600 83 375 1463 20,000 1,829 29,600 No.oflakeendemics 920 75,000 922 No.ofspecies 615 32,900 280 Area (km²) 473 Lake Turkana 1,240 Lake Kivu Lake Tana Altitude(m) 773 Lake Edward Lake Mweru Lake Albert Lake Chad Lake Malawi Lake Victoria Lake Tanganyika Lake assemblage,extendingfrom theMaghreb totheNile, , haverecently beenrecognized asoccurringinthe A. cygnea Unio gibbus c withtwowidespread butdecliningEuropean Anodonta nor Unio U. gibbus U. delphinus (CriticallyEndangered). Moreover, we isanothertypicallyPalaearctic genus, and theyare. Themore traditionalview , et al. U. delphinus A. anatinus asaseparateendemicspecies, 2010;Araujopers.comm.) A. pallaryi U. durieui were evaluatedaspartofthis Unio U. mancus speciesare recognized . Thosetaxaare both , et al. and U. ravoisieri (Endangered). The et al. 2009).However, A. lucasi 2009).Neither tobeanother , and . Dueto U. from thegenusLavigeria. A LakeTanganyika gastropod(demonstratingshellrepair) ornamentation, withshell-formscalledthalassoid,meaning ornamentation, 1994), andsomeendemicspecieshavespectacular in thelittoralandsub-littoralzones(Leloup1953;Brown al. of theMetazoanfaunabeingendemictolake(West of theEastAfricaRiftValley lakes,withapproximately half Lake Tanganyika isviewedasoneoftheoldestanddeepest the pulmonates(Brown 1994). whilst there isamore limiteddiversityandendemismof largest lakes,Tanganyika, Victoria andMalawi(Table. 4.1), In general,theprosobranchs are mostdiverseinthethree distinctive speciestoevolve. been isolatedforsuffi (1994) pointsoutthateventhesesmalllakesmayhave change recognized intheancientlakes,althoughBrown do nothavethesamelevelsofisolationandevolutionary and thisisconfi diversity ofmolluscs(West have long-beenrecognized fortheirhighendemismand 2003).Theendemicprosobranchs are principallyfound rmedhere (Figure 4.2).Othersmallerlakes cient periodsforthepresence of © HEINZH.BÜSCHER et al. 2003;Michel et al. 2004), et CHAPTER 4 | MOLLUSCS 101

. . were were Lanistes Bellamya Coelatura Bellamya , which is sh that are sh that are (Brown 1994) (Brown , are categorized , are , which is assessed as 2011), and the taxonomic Melanoides Nyassunio nyassaensis in Lake Victoria has yet to be has in Lake Victoria et al. Aspatharia divaricata is a thin shelled elegantly shaped ned requiring further investigation ned requiring Eupera crassa Chambardia nyassaensis L. nasutus has a restricted range within the lake, being range has a restricted 2007). Some of the more widespread families widespread 2007). Some of the more was assessed as Near Threatened, whilst was assessed as Near Threatened, and Mutela bourguignati et al. an apple-snail (Ampullaridae) that is restricted to an apple-snail (Ampullaridae) that is restricted dependent on the presence of molluscs as their food dependent on the presence highly specious in Lake A number of genera are source. includes eight Malawi. For example, the family Thiaridae endemic species in the genus that these eight Recent molecular studies have shown shells, do not endemic species, with highly distinctive parthenogenetic clonal lineages genetically and are differ (Genner as also include localized endemic species, such nasutus, Lake Malawi. to 82m. Of the deep-water species found at a depth of 46 mussels endemic to Lake Malawi, two species, freshwater Mutela alata due to impacts of sedimentation, as Vulnerable, hypsiprymna the other endemics such as Least Concern.considered lakes shallower than the other great is Lake Victoria (maximum depth of 80m), has a lower salinity than Lake with extensive and possesses a shoreline Tanganyika, papyrus swamps, open sandy beaches and stony shores is considerable variability within some 1994). There (Brown (1994) suggests these may either be in species, and Brown periods of low lake levels, a status of partial speciation from deﬁ just poorly or are of their genetic distinctness. The African viviparid fauna in the lake included six species in the genus Ongoing investigations by Lange (pers. comm.) should of the true distribution ranges better knowledge provide of populations within the lake as, for example, phthinotropis in the south-west part. Freshwater only found currently mussels of note are and Critically Endangered, (Graf Vulnerable considered status of determined. rigorously between the two lakes (Brown 1994). The shore is mixed, is mixed, The shore 1994). (Brown between the two lakes separated by sandy bays and small shores with rocky is known to be subject to estuaries, and the lake-level rainfall since the Quaternaryseasonal changes in Period. Lake in recognized genera are no endemic Currently, a are Again, there Tanganyika. Malawi, unlike in Lake ﬁ number of lake endemic molluscivorous

), t is shes 2003). which is Moncetia, Nyassunio et al. Spathopsis (and Brazzaea anceyi M. anceyi, In the family Paludomidae, In the family Paludomidae, Chambardia sh, also endemic to the lake, sp X, etc. (West sp X, etc. (West Pseudospatha tanganyicensis cient. Another genus, Richness of bivalves is lower than and Lavigeria ocks in the same way as the cichlid fi (2004) to suggest that the lake molluscs (2004) to suggest that the lake molluscs sp P, sp P, et al. 2003). Various fi 2003). Various which is thought to include 18 species. However, which is thought to include 18 species. However, to be a synonym of , are known from few specimens, and as a result few specimens, and as a result known from , are et al. Lavigeria LakeLake VictoriaLake ChadLake MweruLake TanganyikaLake TurkanaLake Malawi (m) Altitude 1,240 280 773 922 (km²) Area 375 75,000 No. of species 473 No. of lake endemics 20,000 32,900 4,850 7,200 17 29,600 12 11 12 6 species/subspecies 10 1 species/subspecies 8 species 8 4 species/subspecies 1 subspecies 3 species and reported that this species had never been collected that and reported of the study by unaware alive; however he was apparently Kondo (1984). Lake Malawi is less saline and shallower than Lake with a mollusc fauna which is about 60% Tanganyika, taxa shared endemic to the lake and with no gastropod Table 4.2. Freshwater bivalve fauna of the largest African lakes. largest African of the bivalve fauna 4.2. Freshwater Table These species vary in their threatened status, depending These species vary in their threatened sedimentation on their range and the localized impacts of would benefi These species groups along the lake shore. as more once the taxonomy has been revised, review from if their ranges are species may be found to be threatened thought. The numerous than currently restricted more variable shapes, species, with extremely prosobranch led Michel formed species fl (West are dependent on the presence of these molluscs as their dependent on the presence are food source. bivalves: Freshwater African Lakes (see Table in the largest that of gastropods mussels genera of African freshwater three are 4.2) There and endemic to Lake Tanganyika. each monotypic that are Grandidieria burtoni are assessed as Least Concern, assessed as are but and is listed to Lake Tanganyika also endemic apparently Mandahl-Barth (1988) considered as Near Threatened. Moncetia ‘sea-like’, as there is a marked convergence with marine is a marked convergence ‘sea-like’, as there found molluscs are (1994) suggests that few shells. Brown to the concentration of hydrogen below 125m depth, due sulphide. prosobranchs: Freshwater and 20 genera belong to the subfamily Hauttecoeuriinae, The most to Lake Tanganyika. restricted 19 of these are dwelling genus speciose genus is the non-thalassoid rock Lavigeria, of these although a diagnostic key does exist, a number being indicated species have not yet been fully described, by Other mussels, such as Vulnerable. considered ujijiensis their status is Data Defi by a single species is represented 102 CHAPTER 4 | MOLLUSCS View fromnearKigali,Rwanda. Lakesregion oftheGreat extirpated (Van Damme prosobranchs are assessed asthreatened orare locally Damme in thelimestonemountainousregions (Ghamizi1998;Van in theMaghreb region isfoundinspringsfedbyaquifers – 49%(Strong (around 105species –18%)andAustralia(252species species –69%(Seddonpers. comm.)),NorthAmerica (69 species–11%),incontrastwithEurope (around 610 spring-snails formalowerproportion oftheendemicfauna Hydrobiidae. InAfrica,especiallytheMaghreb region, and largely comprisesthespring-snailsfrom thefamily many otherpartsoftheworld,foundinartesianbasins, The highestdiversityoffreshwater molluscsis,asfor 4.1.3 Molluscandiversityinartesianbasins which isassessedasEndangered. regarded asVulnerable, and Species ofnoteare well astwoendemicmonotypicfreshwater musselgenera. shallow lake.There are eightendemicGastropoda taxa,as Brown (1994)commentedontheunusualrichnessofthis (1927) commentedonthevulnerabilityoffauna,and papyrus swampsalongthefringes.PilsbryandBequaert (15m depth(Brown 1994)),withextensivemarshesand drains intothesouthedgeoflake,whichisveryshallow also present inthelowerreaches oftheLuapulaRiverwhich these speciesare restricted tothelake,whereas othersare Zambia, hasseveralendemicspeciesofmollusc.Some Lake Mweru,alakeintheupperpartofCongoBasin et al. 2010).More than75%oftheMaghrebian et al. 2008)).Themajorityofthisdiversity Mweruella mweruensis et al. Prisodontopsis aviculaeformis 2010).By contrast,there are , whichis © IUCN/INTUBOEDHIHARTONO , (Arconada andRamos2003;Benke glacial limit(e.g.,theBalkansandIberianpeninsula radiations are foundinthelimestoneregions southofthe endemic genus.InneighbouringEurope, thegreatest One ofthese, origin oftheseradiationsandthelacklimestoneregions. fauna southoftheSahara,possiblyarefl proportionately fewendemichydrobiids intheAfrotropical coastline. Thesalinelagoons havesomevery restricted range saltmarshes and mangroves thatfringe large partsofthe range-restricted speciesare foundinthesalinelagoons, show littlespatialvariation.However, someofthemore Molluscs ofthecoastalhabitats ofAfrotropical Africa 4.1.4 be threatened. tolerant ofdrying-outand,assuch,are notconsidered to in thesehabitatsare typicallymore widespread species The freshwater bivalvesandmostofthepulmonatesfound within thesesystems. and, assuch,there maybefurtherunknowndiversity due tothediffi with artesianwatersthatremain tobesurveyed,largely 20 genera.There are, however, large partsoftheregion particular inAlgeria,couldraisethenumbertomore than from theregion, butitislikelythatfurtherinvestigation,in of subterraneanhydrobids. Atleast13generaare known Africa, theMaghreb isacentre ofevolutionandradiation saltmarshes andmangroves Molluscan diversityinsalinelagoons, Lobogenes culty ofsamplinginterstitialgroundwaters , is a central and southern African , isacentralandsouthern et al. 2009)). In northern 2009)).Innorthern ection ofthe CHAPTER 4 | MOLLUSCS 103

Gabbiella ; these taxa western Uganda crater tropicus toroensis cient below 20m depth and, in

which is distributed widely in the widely in which is distributed

, (EN). In the Bulinus and Molluscan diversity in crater lakes Molluscan diversity Lake Awing, one of the crater lakes in Cameroon. © KLAAS-DOUWE DIJKSTRA cient. Mytilopsis africanus Mytilopsis western Africa, extending from Senegal to the Congo. Congo. to the Senegal western from Africa, extending Dreissenidae of the representative Afrotropical The sole is in many parts of Africa, including present Crater lakes are Ethiopia and eastern Africa. These small, Cameroon, as usually species-poor, but deep lakes are sheltered, often oxygen deﬁ they are Some species too saline for molluscs. some cases, are to colonise these lakes, probably managed have, however, isolation and their relative though transportation by birds, species. The may have driven their evolution into new crater lakes have a richer fauna than might Cameroon species, be expected, including one possibly endemic Bulinus camerunensis estuarine/brackish waters of coastal western Africa, from western waters of coastal estuarine/brackish Africa, from Most of these species the mouth of the Congo. Senegal to the as Least Concern, assessed although some, where are as Data categorized are uncertain, data were distributional Deﬁ 4.1.5 recorded bivalves were and three lakes eight gastropods by Mandahl-Barth, including two endemics, kichwambae further investigations, using molecular systematics, require 1994). to establish their status (Brown

nd which are which are in the Oued Abid 2009a), so the 2009a), so found in southern found Saulea, et al. and Tomichia are sensitive to changes in to changes sensitive are , of the family Cyrenoididae, , of the family Cyrenoididae, Tomichia Afropomus Margaritifera marocana Margaritifera . Most of the African littorinids are not not littorinids are . Most of the African Cyrenoida Littoraria occur in coastal western Africa (plus four species majority have been assessed as threatened. been assessed as majority have be more species tend to the mangrove By contrast, found in suitable habitats throughout and are widespread, example, the family Littorinidae is the continent. For in Africa by a few represented a cosmopolitan family species of in some regions, However, to be threatened. considered swamps of westernsuch as the coastal you fi Africa, the “living fossils”, their habitats caused by pollution and interference with the and interference caused by pollution their habitats (Kristensen regime seasonal wet-dry species, such as the species of the species such as species, of All species Africa. A team of Moroccan and Spanish malacologists aided by local youngsters collecting (Morocco). After tissue samples were taken, the animals of this last viable population of the species were placed back in the river. © MOHAMMED GHAMIZI are found in brackish/estuarine habitats in tropical found in brackish/estuarine habitats in tropical are of unknown distribution that were not evaluated; one of unknown distribution that were invalid). species that was evaluated is now considered species of Two monotypic, are considered to be rare, and are threatened. threatened. and are to be rare, considered monotypic, are typically found in brackish families of bivalve are Three mainly comprised of water habitats, and these families are 19 species marine species. Recent checklists have listed western tropical Africa (Daget of the family Donacide from 14 of these have only 1998; Bogan 2010). However, of species Ten been evaluated for the IUCN Red List. Egeria 104 CHAPTER 4 | MOLLUSCS Table 4.3.ThenumberofAfricanmolluscspeciesineachIUCNRedListCategory, byregion. in 1994.Thissurveyusedthefi of thefreshwater gastropods undertakenby DavidBrown Red ListofThreatened Species The fi 4.2 continental Africa,inthegastropod groups: a)theprosobranchs; b)thepulmonates;andc)bivalves. Figure 4.3.Theproportion (%)offreshwater molluscspeciesineachregional IUCNRedListCategoryinmainland 4.1.6 clams isolation. AlthoughmostspeciesoftheAfro-tropical pea- some endemicspecies,possiblyalsoasaresult oftheir As forthecraterlakes,theseare isolated,andsupport (amountainlakeorpool)athighaltitude. have smalltarns The highmountainareasAfricaandEthiopia ineastern and increased frequency ofdrought. potential changesinlakelevelsduetoclimatevariability species forsportfi lakes are threatened bytheintroduction ofnon-nativefi Threatened ( al. Africa(Waruihigh-elevation montanelakesinEastern a fewlakesinEthiopia.Two more speciesoccurringin Critically Endangered, havingadistributionlimitedto some species,suchas (sixspecies)orDataDefi Concern precautionary measure notincluded inthepan-Africalistingsforendemic species. *Some subspecies regarded asspecies inregional treatments, andundescribedtaxa includedinregional treatments, are as a otenArc 5 5 17% 19% 16% 24% 38% 45% 31% 12% 9% 22% %DataDefi 155 cient 159 95 %Threatened 122 228* Pan-Africa NoofSpecies Africa Northern Central Africa Western Africa Africa Southern Africa Eastern Region 2001)are assessedasVulnerable ( rst speciesfrom theregion tobeplacedontheIUCN molluscs Conservation statusoffreshwater Molluscan diversityinmontanelakes ( Pisidium P. montigenum spp.) are assessedasbeingofLeast shing, aswellbytheirvulnerabilityto P. ethiopicum, ). Speciesinthesemontane rstsetofquantitativeIUCN

came from amajorreview 2*2%28% 29% 624* cient (sixspecies), P. artifex are considered ) andNear sh et Defi proportion ofthreatened speciesandtheproportion ofData There isconsiderablevariabilitybetweentheregions inthe ongoing globalassessmentofallfreshwater molluscs. ranges. Theseregional assessmentswillcontributetoan to determinetheirstatuswithintheAfricanpartof Africa, regional assessmentshavealsobeencompleted List. Forthosespecieswithdistributionsextendingbeyond for allendemicspeciesare addedtotheglobalIUCNRed equivalent totheirglobalstatus,suchthatassessments continental Africa,theirregional IUCNRedListstatusis regional scale(IUCN2001).Forthosespeciesendemicto IUCN RedListCategoriesandCriteria:Version 3.1ona of freshwater molluscwasassessedbyapplyingthe this project, theconservationstatusofeachspecies status oftheregion’s freshwater molluscanfauna.In provides acomprehensive overviewoftheconservation This IUCNpan-Africanproject isthefi biodiversity forthecontinent. could notprovide anoverviewofthestatusfreshwater species,andsothefi or LeastConcern did notincludeassessmentsforanyoftheDataDefi review, publishedbyIUCN(BaillieandGroombridge 1996) Rift Valley Africa.However, lakesandwestern Brown’s mainly thosefoundintheregional hotspotsintheAfrican criteria (version2.3)andresulted inthelistingof68species, cient species.This,inpart,refl ects thelevelofrecent grs obtained gures rst studythat cient CHAPTER 4 | MOLLUSCS 105

et al. the majority of lling. There is, however, an ongoing is, however, lling. There in the Lower Congo Rapids and Malebo

. The Lake Chad catchment is shown to include a Sites highlighted as having the most threatened species Sites highlighted as having the most threatened the East African Lakes (Lake Victoria, include the Maghreb, and Lake Malawi) and the Congo Basin Lake Tanganyika 4.4) (Figure number of threatened species, but these require taxonomic species, but these require number of threatened have not been highlighted as requiring and, as such, review conservation actions. In the Congo Basin, Pool were assessed as threatened, representing a quarter representing as threatened, assessed Pool were (Graf of the total molluscan richness in those areas species living range-restricted 2011). Most of these were within specialised habitats, such as rapids. Pilsbry and Bequaert (1927) commented on the vulnerability of the fauna of the Lake Mweru in central Africa to its gradual infi endemic species eld survey, the eld survey, ect regional differences differences ect regional These factors should be These factors should be taken into consideration when interpreting the species taken into consideration when interpreting distribution maps. taxonomic research, the amount of recent fi the amount of recent taxonomic research, application of molecular systematic, and the number of working on each local scientists and overseas researchers most the are In general, the prosobranchs taxonomic group. by the followed molluscs group, of the freshwater threatened 4.3). 4.3; Figure bivalves and then the pulmonates (Table species distribution patterns 4.2.1 Threatened variations in the distribution patterns regional Apparent in part refl species of threatened levels of survey in the status of taxonomic research, endemic species, work, and numbers of range restricted threatened. often which are Figure 4.4. The distribution of threatened freshwater mollusc species across mainland continental Africa. continental mainland across species mollusc freshwater of threatened distribution 4.4. The Figure sub-catchment. per river/lake of species = number richness Species 106 CHAPTER 4 | MOLLUSCS Cape Province, SouthAfrica. neartheEastern only knownfrom onelocation,astream Tomichia cawstoni species inthegenus precautionary approach, suchthatthethree endemic considered threatened. Here, wehavefollowedthe in thelowerreaches oftheLuapulaRiverthatwere not both theendemicspeciesandthosealsopresent debate astowhethertherateofinfi ( Endangered ( one assessedasExtinct( the 1996IUCNRedList(BaillieandGroombridge 1996), occurring inSouthAfrica,three specieswere includedin changed thetotalnumberofspeciesand the previous assessmentin1996,andsomeofthesehave There havebeenseveralmajorchangesintaxonomysince 4.2.1.1 Changessincethe1996assessment species are relatively widespread. al. other groups ofthreatened freshwater taxa(Darwall The Zambeziheadwaterswere considered importantfor are monitored closely. its limitedoccurrence, itisstillimportantthatpopulations assessed asNearThreatened. Taking intoconsideration the springsthatthesesnailsinhabit,anditistherefore threatened, asthere are noknown plausiblethreats to rogersi Tomichia recent review ofthepresent status ofpopulationsall vulnerable todestructionbyman(Brown 1994).Thismore Herbert (1998)notedthatofthesevenspecies temporary salinepanstoperennial freshwater streams. are foundinthecoastalregions inhabitatsrangingfrom Africa,someofthemoreIn southern unusualendemics aviculaeformis mweruensis are alsoendemictoLakeMweru,ofwhich Endangered. Two monotypicfreshwater musselgenera Tomichia 2009),butlesssoformolluscs,asthemajorityof hasarestricted range,itcannotbeassessedas speciesinSouthAfricanotedthatalthough

rogersi isregarded asVulnerable and Tomichia natalensis isassessedasEndangered. ), asthehabitatsofallwere considered , a Critically Endangered species currently speciescurrently , aCriticallyEndangered Bellamya Tomichia © DAIHERBERT are assessedasCritically ), andoneEndangered

cawstoni ll islikelytoimpact Prisodontopsis ), oneCritically Mweruella Tomichia et T. T.

the northern African region,the northern aswell southandeast throughout therest ofAfrica. Thisistrueformostof Restricted range speciesare found onlyexceptionally Atlas, andMoyenAtlasmountain rangesintheMaghreb. Congo; andtheisolatedHaut Atlas, AtlasSaharien,Anti of thevast,diversiﬁ presence ofthelarge riftsystem,and lakes intheeastern AfricaandinCameroon; partofwestern the the southern of somethesmallercoastaldrainages,particularlyin partly duetoanumberofreasons: thegreat relative age gastropods, mostofwhichare prosobranchs. Thisis Africa isacontinentwithmanyrestricted rangefreshwater 4.2.2 Restrictedrangespecies criteria themselves. information, andchangestothethresholds oftheRedList status ofthespeciesbutrathertaxonomicchanges,new made byBrown (1996)reﬂ observed changesincategoryfrom the1996assessments data shouldbeinterpreted. Inconclusion,fewofthe are conducted,including,forexample,thewaythatrange have ledtoanumberofchangesinthewayassessments Categories andCriteria(version3.1)theguidelines effort mentionedabove,thelatestversionofIUCN In additiontothetaxonomicchangesandincreased survey previously knownfaunaandthenewlydescribedspecies. has increased ourknowledgeofthedistribution along withtheGEFLakeTanganyika BiodiversityProject, research andsurveyworkbyWest the LakeTanganyika gastropods resulting from recent There havealsobeennewspeciesdescriptionswithin sources, andhenceare nowconsidered tobethreatened. as rangerestricted, withthreats from exploitationofwater been spiltintomore than13species,someare regarded therefore however, LeastConcern; nowthe specieshas species, descendants are exactcopiesofthemothers.Asasingle the different morphshavebeenverystable,andthattheir al. colour butvaryverylittleingeneticmake-up.Heller clonal lineagesthatdiffer spectacularlyinshellform and single species,arguing thatasexualreproduction creates as wasBrown’s (1994)opinion.Brown considered thema rather thanbelongingtoasinglespecies, currently considered asdistinctspecies(Heller some ofwhichare CriticallyEndangered. Theseare Maghreb, where itisrepresented byabout13species, 1988). Onegenusinthisfamily, Thiaridae onthebasisofanatomicalcharacters(Houbrick The familyMelanopsidaewassplitfrom the family species conservationstatus. range restricted species,withconsequentimpacts to (2005)reviewed thegeologicalhistoryandfoundthat M. praemorsa edandoldhydrographic systemofthe was considered widespread and ect genuinechangesinthe Melanopsis, et al. (2003).Thiswork, M. praemorsa occursinthe et al. 2005) et ,

CHAPTER 4 | MOLLUSCS 107 some of which ood events at outlets Tomichia, ned to small stretches of the coast of Natal and ned to small stretches (found on the coast of Natal) and conﬁ are the Cape Province. ranges, sometimes of Species with the most restricted hydrobids the Maghrebian are metres, only a few square one or two springs or wells. Although found in that are extensive, they may be more their true range underground as basis, on a precautionary threatened assessed as are the extent of their range in artesian waters is uncertain; to the wells, whereas some species may be truly restricted only seen in ﬂ others which are widely in the groundwater more actually distributed are supplying the outlets.

Septaria uctuations during the of the Atlas range, i.e., Mauritania, Mali, Libya, central and southern Algeria and Egypt, the whole of the north- eastern Sudan, Ethiopia, Djibouti African Region (Eritrea, and Somalia), with the exception of a couple of endemic species in south Somalia and south Ethiopia. The same is also true for the northern part of the western African basins of the Niger and Senegal that i.e., the region, some drying events during the Pleistocene severe suffered 20,000 years ago. In southern climate fl Africa, severe inhospitable for Afrotropical Pleistocene made this region range species no restricted are there so prosobranchs, with the exception of species within the genera Figure 4.5. The distribution of restricted range mollusc species in mainland continental Africa, showing Africa, showing continental in mainland species range mollusc of restricted distribution 4.5. The Figure river/ per of species = number richness Species Lakes. the Rift Valley and Maghreb in the of diversity hotspots lake sub-catchment. 108 CHAPTER 4 | MOLLUSCS River nearSoapiti,Liberia,and monotypic genera(e.g., Somalia toMozambique,anumberofspeciesandeven as wellalongthecoastofIndianOceanfrom southern coast, from Côted’Ivoire totheRepublic oftheCongo, In smallriverbasinsandmarshesalongAfrica’s Atlantic is notthe easiestplace tosample), manyare habitat than thetypelocality (thestretch of rapidsnearMatadi easily overlooked. Whiletheirtruerangemust belarger Hydrobiidae, Assimineidae andBithyniidae,whichare the caseforsmallsizedspecies belongingtothefamilies Congo) are onlyknown from thetypelocality. Thisisonly Congo RiveratAngo-Angonear Matadi,Republicofthe status. Speciesrichness=numberofspeciesperriver/lakesub-catchment. research, fi of speciesinthecentresAfricarequire ofdiversityintheAfrican lakesandnorthern additionaltaxonomic Figure 4.6.Distributionofthefreshwater molluscsassessedasDataDefi eld survey, andinformationonrelevant threats inorder tomakeanassessmentoftheirconservation Soapitia dageti Valvatorbis mauritii from theKonkouré inthe deep, meromictic lakeslike LakesTanganyika andMalawi, over whichitcan impactwithinashortperiod oftime.In exercise, asthisdepends onthelevelofthreat and thearea rift lake,whileothersdidnot.Admittedly, itisnotaneasy threats likelytospread throughout andimpactawhole not applythesamecriteriaforall lakes,someconsidering (Darwall In thereport Africanregion onthemolluscsof the eastern Finally, there are theendemicsoflarge RiftValley lakes. also beenassessedasthreatened. such asstonesinrapids.Thesespecieshave,therefore, specialists confi et al. 2005),theoriginalassessorsapparently did cient, showingthatlargenumbers ned toaspecifi c habitatoflimitedextent, CHAPTER 4 | MOLLUSCS 109

ed cient Burnupia cient due cient due and Ferrissia ciency in molluscs are ciency in molluscs are ed as Data Defi cient species cient cient. This contrasts with western Africa, 4.2.3 Data Defi for Data Defi The two main reasons knowledge. taxonomic uncertainty and poor geographic of Data Defi with the highest proportion The regions northern Lakes, and species are Africa, the east African western 4.6). Africa (Figure In southern to a number Africa, taxonomic issues relating of small limpet species in the genera are the main reasons behind these species being classifi the main reasons are as Data Defi as survey data available and, is little recent there where identifi were species such, more to lack of recent data on distributions (Figure 4.6). In the data on distributions (Figure to lack of recent eastern, western and southern knowledge African regions, due to surveys conducted in is, in general, reasonable funded by the World the second half of the 20th century, snails Health Authority (WHO) to study African freshwater the transmission of schistosomiasis and in that play a role other parasites. As no schistosomiasis occurs in northern was not incorporated in the research Africa, this region subsequently are campaigns funded by WHO, and there a higher number of species listed as Data Defi to a lack of distributional data. In northern Africa, the high molluscs level of taxonomic disarray within the freshwater a consequence of many species being described is largely “Ecole Nouvelle”, by members of the 19th century French © DANIEL GRAF AND KEVIN CUMMINGS cient or Near ) are considered considered ) are , which is reported as only , which is reported L. solidus of Lake Albert, the status , which has only been recovered , which has only been recovered and ned to a small oxygen-rich stretch 2009b). Such species have been et al. Bellamya phthinotropis Lanistes nyassanus the benthic fauna is confi and upper close to the coast (upper 80m in Lake Malawi, the anoxic or hypoxic In 100 to 200m in Lake Tanganyika). few exceptions, Barring a is no macro-life. zones there as endemics have been assessed all Lake Tanganyika (Least Concern,not threatened Data Defi occur in a whilst range-restricted, as many, Threatened) for the threshold number of locations than meets greater of the given the localized nature category, a threatened some Lake sublittoral zone. In contrast, to the lake’s threats the sublittoral Malawi endemics, though found throughout (e.g., but poor quality of data available at the as Endangered, the assessments to time of assessment may require Ongoing investigations leading to a better be revised. distribution ranges of populations knowledge of the real in these lakes will soon be available. The same is true for information current of other lakes where the prosobranchs distributions within lakes, shows species to have restricted such as For other endemics, such as assessed as threatened. Gabbiella humerosa alberti being found in the south-west part of Lake Victoria, and being found in the south-west part of Lake Victoria, Gabbiella neothaumiformis in the south-west part of Lake area a very restricted from Chad (Kristensen during the next may need re-evaluation of Endangered information becomes available. assessment as more Diving for mussels in the Upper Chambeshi River, Upper Congo. River, Diving for mussels in the Upper Chambeshi 110 CHAPTER 4 | MOLLUSCS Table 4.4.Extinctmolluscspecies inAfrica. activities. and theimpactsofthreateningdistributions patterns extensive fi survey workhasbeenconducted. Theseregions require with alonghistoryofconfl Africa,Angolaand Mozambique, all countries insouthern only afewmalacologistsventured intotheregion. Similarly, available forSomaliasince1930, andevenbefore 1930 number ofareas. Forexample,there are virtuallynodata unrest havealsolimitedtheavailableknowledgefora Diffi Defi from theEritrean islandsonly, are alsorecorded asData many mangrove species,someofwhichwere described made tore-examine itsincethelate19thcentury. Assuch, fauna isquiterich,butapparently noefforts havebeen in theregion. However, bycontrast,thebrackishwater freshwater, andhencethere hasbeenlittlesurveyeffort have received littleattention.Eritrea hasvirtuallyno Africa,theregionsIn north-eastern ofEritrea andSomalia also includedintheWHOcampaigns. Defi work. InEgypt,EthiopiaandSudanthenumberofData has emerged whoare startingtodosomeoutstanding changing asanewgenerationofMaghrebian malacologists populations sincethatperiod.Thissituationisgradually and westilllacksuffi 1920, there remains doubtastotheirtaxonomicvalidity, for quiteanumberofspeciesdescribedbetween Few recent taxonomicrevisions havebeenmade,sothat, where allvariantswere givenadifferent speciesname. aiyGnsSeisIUCNStatus Species Genus Bithyniidae Family Bithyniidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae culties inaccessingsomecountriesduetopolitical cient. cient speciesisrelatively low, asthesecountrieswere eld survey inorder tobetterunderstand species cient informationonthestatusoftheir ihnatentaculata Bithynia ihnaleachii Bithynia yhnlalimnopsis Bythinella yhnlamauritanica Bythinella yhnlamicrocochlia Bythinella yhnlapunica Bythinella yrbagracilis Hydrobia ecraletourneuxiana Mercuria suancl doumeti Pseudamnicola suancl globulina Pseudamnicola suancl barratei Pseudamnicola suancl desertorum Pseudamnicola suancl oudrefi ca Pseudamnicola suancl ragia Pseudamnicola suancl latasteana Pseudamnicola suancl singularis Pseudamnicola icts, havelarge areas where little ca 1850- fi from South Africa,maybealready Extinct,butmore extensive and kadeii that someofthecentralAfrican species, suchas are noprosobranchs listedasExtinct,thoughitisprobable localities inMediterraneanAfrica. Intherest ofAfricathere been lostfromedges oftheirrangesinafew thesouthern still foundthroughout Europe andAsiaMinor thathave Africa. Thesearenorthern widespread Palaearctic species The RegionallyExtinctgastropods are also restricted to two orthree timesthenumbercitedabove. of extirpatedspecieswillneverbeknown;theycould possible toreview thetaxonomysuchthattruenumber lost ordisturbedatmanylocations,soitwasnolonger the watersources andfreshwater springs were already Boeters (1976)triedtore-investigate thesitesin1970s lumped inthe19thcentury. Bythetimeresearchers suchas higher, asthissubterraneanmalacofaunawastaxonomically out. Thenumberofextirpatedspeciesmaybeconsiderably underlying aquifers,whichhasledtomanyspringsdrying and theover-abstraction ofwaterfrom thespringsand species havesuffered from acombination ofwaterpollution only oneortwosprings,wellsthermalsources. These small hydrobid snailsthatare range-restricted, occurringin be Extinct(Table 4.4).Thisregion hasahighproportion of Algeria andTunisia), where 14speciesare considered to lie intheMaghreb regionAfrica(Morocco, ofnorth-western in AfricasouthoftheSahara,butmostconfi There are afewpossibleextinctionsorregional extinctions 4.2.4 Extinctionpatterns eld surveysare required before thiscan be confi Tomichia cawstoni (here listedasCritically Endangered) from Cameroon Regionally Extinct Regionally Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct Extinct (alsolistedasCritically Endangered) rmed extinctions rmed. rmed. Potadoma Potadoma CHAPTER 4 | MOLLUSCS 111 (2003) on the VU et al. 2010). In Lake et al. species along the lake Probably LC Probably threats threats threats threats Lavigeria (2004) and West (2004) and West rmed as valid species may be et al. subsp. nov. ‘boulanouari’ subsp. nov. subsp. nov. ‘taramtensis’subsp. nov. CR ed as threatened by sedimentation and pollution ed as threatened Tanganyika, Michel Tanganyika, many new possible report appear to be quite widespread, Many of these margins. be probably and hence, when formally described, will at least three, assessed as Least Concern. However, taxa have a limited of these undescribed possibly four, range, and once confi classifi adjacent urban areas. from are found have been severely depleted through habitat depleted through found have been severely are habitat and through of water, loss due to over-abstraction pollution and saline intrusions from degradation resulting Damme (Van into their groundwaters (2003) p. 46 DD LC Probably (2003) p. 49 (2003) p. 56 (2003) p. 56-7 DD DD DD Uncertain range extent; DD? LC Probably (2003) p. 43 DD Uncertain range extent and (2003) p. 52 DD LC Probably (2003) p. 50 (2003) p. 54 (2003) p. 41-2 DD DD DD LC Probably LC Probably Limited range, but uncertain (2003) p. 42 DD Limited range, but uncertain (2003) p. 52 DD LC Probably (2003) p. 53-4 DD LC Probably (2003) p. 55 (2003) p. 51 DD DD LC Probably LC Probably (2003) p. 45-6 DD LC Probably (2003) p. 47 DD LC Probably et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. andraea sp. nov. ‘wiwanensis’sp. nov. VU sp. nov. ‘mirlheftensis’sp. nov. EN sp. nov. ‘haasei’sp. nov. EN sp. nov. ‘aghbalensis’sp. nov. EN sp. nov. ‘salahi’sp. nov. EN sp. nov. ‘boulali’sp. nov. EN sp. nov. ‘meskiensis’sp. nov. EN sp. nov. ‘makhfamanensis’sp. nov. CR sp. nov. ‘kerdouensis’sp. nov. CR sp. nov. ‘valai’sp. nov. CR sp. nov. ‘tiznitensis’sp. nov. CR sp. nov. ‘ramdanii’sp. nov. CR molluscs endemic to Africa. cient freshwater sp. J West sp. F West sp. T West sp. S West sp. P West sp. K West sp. R West sp. B West sp. D West sp. N West sp. H West sp. C West sp. Q West sp. O West sp. G West sp. M West Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Lavigeria Heideella Belgrandia Mercuria Horatia Horatia Heideella Heideella Giustia Semisalsa aponensis Heideella Heideella Heideella Bythinella Belgrandiella Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Paludomidae Paludomidae Hydrobiidae Paludomidae Hydrobiidae Paludomidae Paludomidae Hydrobiidae Paludomidae Paludomidae Hydrobiidae Paludomidae Paludomidae Paludomidae Paludomidae Hydrobiidae Paludomidae Hydrobiidae Paludomidae Hydrobiidae Paludomidae Paludomidae Hydrobiidae FamilyPaludomidae Genus Species Source RL Category Possible Status FamilyHydrobiidae Genus Species Subspecies RL Category These are all species that have been listed in books or PhD. theses but are yet to be formally described, so are not yet placed not yet yet to be formally described, so are all species that have been listed in books or PhD. theses but are These are IUCN Red List. 4.2.5 Undescribed species species in northern some recognized Africa are There that have not yet formally and Lake Tanganyika with clearly threatened been described which are formal 4.5). These species require extinction (Table to be included in the IUCN Red descriptions in order described also List. Other species that have yet to be investigated exist in Africa, and have yet to be fully 4.6). (Table species known from range-restricted Most of these are in which they northern the water sources Africa, where Table 4.5. Undescribed threatened freshwater molluscs endemic to Africa. to endemic molluscs freshwater threatened 4.5. Undescribed Table Table 4.6. Undescribed Data Defi Table 112 CHAPTER 4 | MOLLUSCS increased frequency ofdrought. by “naturaldisasters”,whichinthiscaserefers largely tothe of threatened species.Finally, somespeciesare threatened are onlyrecorded asasignifi present andhavehadanimpactonsomespecies,butthey on nativespeciesofmollusc.Invasivemolluscare invasive speciesdonotappeartohavehadanotableimpact and Brown 1999).Incontrasttoothertaxa,suchasfi gastropod species,whichare vectorsofbilharzia(Kristiansen of Burkina Faso. Pollution ofasmall riverclosetoOuagadougou, thecapital activity inthecatchment,useofmolluscicides,and due toincreased levelsofsedimentationfrom agricultural Africa,threatsIn eastern from declininghabitatqualityare fl combination ofwaterpollution,sedimentation,altered river from SierraLeone,Côted’Ivoire andGuinea,through the This impactsanumberofrestricted rangeendemicspecies damming riverfl in crop plantations,expansionofbauxiteminingand primarily asaresult oftheextensiveusepesticides Africa,thehabitatqualityinriversisdeclining In western from thedirect impacts from region toregion. Anumberofspeciesare alsosuffering threatened species,butagain,thesources ofthepollutionvary of sources isanothermajorthreat, whichimpacts63%of regions, asexplainedbelow. Water pollution from arange throughout Africa,thecausesofhabitatdeclinevarybetween the consequences.Althoughthisproblem iswidespread deforestation, with58%ofthreatened speciessuffering expansion, infrastructure development(e.g.,dams) and degradation, through multipleactionssuchasagricultural 4.7). Themostsignifi threats thatcombinetoleaddecliningpopulations (Figure in Africa.Inthemajorityofcasesthere isusuallyaseries of There are multiplesources ofthreats tofreshwater molluscs ofthreat4.3.1 Regionalpatterns 4.3 Mainthreats species facingthesecombinedthreats. outambensis ow regimes andincreased watertemperature. (CR)and ows forhydro-electric powergeneration. © TIMOMORITZ cant ofthesethreats ishabitatlossor Soapitia dageti of molluscicides employed to reduce of molluscicidesemployedtoreduce cant threat forfewerthan5% (CR)are twonotable Sierraia shes, shes, © TIMOMORITZ Newly builtdam forirrigation,south-westBurkina Faso. the nativespecies. could havelong-termimpactsonthepopulationlevelsof molluscivores oromnivores, whosepredation onmolluscs stocked withintroduced fi (Van Damme Nile Deltawithsubsequentimpactsonthelocalecology the reduced fl habitats inaquifersunsuitableformanyspecies.InEgypt, systems, changingthewaterqualityandleaving in theintrusionofseawaterintokarsticunderground endemics, waterextractionfrom theaquifershasresulted the Maghreb, withitsnumerous underground hydrobid expansion andagriculturalexploitationare highest.In disappearing, particularlyinthelowlands,where urban Damme ecological carryingcapacityoftheenvironment (Van where demographicgrowth hasbyfaroverreached the African region whichisrelatively green andfertile, clearly observedintheMediterraneanpartofnorthern Damme use ofdetergents, chemicalfertilizersandpesticides(Van signifi that are sparselypopulated,waterpollutionisoftenquite and untreated (Van Damme the subsequentreturnofwatersthatare heavilypolluted waters fordomestic,industrialandagriculturaluse, of the Africa,habitatqualityhasdeclinedasaresultIn northern molluscicides (Kristiansen Africa,anduseof mining waste,duneinsouthern from domesticandindustrialsources, sedimentationfrom over-abstraction ofwaterforirrigation,pollution with damconstructionforhydro-electric powergeneration, Africa,thedeclineinhabitatqualityisassociated In southern 2005). LakeVictoria andthe benthic organisms inLakeTanganyika (McIntyre of increased sedimentationisrecognised toimpactmost on theimpactofsedimentationmolluscs, Nile Perch. Although,asyet,there are nospecifi impact ofinvasivespeciessuchaswaterhyacinthand cant, inmanycasesasaresult ofthenowcommon over-abstraction of surface andunderground et al. et al. ow oftheNilehasresulted inaretreat ofthe et al. 2010).Themostdramaticeffects canbe 2010).Inthisarea, speciesare rapidly 2010). sh species,manyofwhichare et al. et al. Ethiopian lakeshavebeen 2009a). 2010).Eveninregions c studies c et al.

CHAPTER 4 | MOLLUSCS 113 uctuations in water level plus cial construction. For example, sh-host movement, potentially (2010) reported that the gastropod fauna the gastropod that (2010) reported et al. Cut off upstream populations of molluscs from populations of molluscs from upstream Cut off populations; downstream habitats and Lead to a loss of the underlying riverine their fauna; part of of the dam, where Cause changes downstream the river is impacted by ﬂ and water temperatures, changing water-chemistry changing potentially impacting life-cycles through success; and patterns and reproductive reproductive Cause a loss of ﬁ success. mussel reproductive changing freshwater

uniformly negative nor positive (Seddon 2000). Dams and uniformly negative nor positive (Seddon 2000). form insurmountable barriers that will: their reservoirs a) b) c) d) examples of benefi are There Damme Van of the Aswan Dam seems to have of the Nile downstream this might alternatively of although improved, be an effect cessation in the use of huge amounts of molluscicides, as employed during the anti-bilharzia campaigns. According populations have also prosobranch (1994), large to Brown developed in the man-made Lake Kariba, but that seems to be an exception. In contrast, after the construction of in the number was an increase in Ghana, there Lake Volga loss of bilharzia-carrying species, and a corresponding of the native pulmonate species (Seddon 2000), with an overall decline in species diversity and species richness. and North America indicate Europe In general, data from biodiversity loss following dam construction, with the sh in existing sh ponds along riverbeds. ow alteration of clear streams shaded by shaded streams ow alteration of clear sh farming. After clearing gallery forests, areas of swamp areas sh farming. After clearing gallery forests, Figure 4.7. Principal threats to freshwater molluscs in mainland continental Africa. continental mainland in molluscs to freshwater threats 4.7. Principal Figure Such actions have changed the ecology, which is now no Such actions have changed the ecology, longer suitable for many of the endemic prosobranchs. 4.3.2 Dams In North America, dam construction during the early declines in the native molluscan 1900s led to severe faunas (Bogan 1993, 2008), and similar patterns now are beginning to be seen on other continents. In Africa, large of for many constructed or proposed already dams are the river catchments, to generate electricity for industrial already and domestic use. The impacts of these dams are being seen for a number of species, and the scale of over the next this development is set to expand greatly few years (Economic Commission for Africa 2003). Dam molluscs in different construction can impact freshwater ways, depending on the life-history strategy of the animal, making the impact of the construction of dams neither In tropical western is major threat In tropical and central Africa, the the siltation and fl red in open waters coloured often resulting gallery forest, laterite muds during the rainy season. Pollution by from as is in this region, raw sewage is another important threat fi too wet for farming and too often formed that are are for raising cattle. As a consequence, and in an dangerous has introduced to diversify local livelihoods, the FAO effort material for stocking fi methods and provided lakes as well as in man-made fi 114 CHAPTER 4 | MOLLUSCS tuberculatus exception ofopportunisticspeciessuchas Africa 2009b). Rawsewageisalsoaproblem inmanypartsof detergents, chemicalfertilizersandpesticides. often quitedramaticbecauseof thenowcommonuseof in regions thatare sparselypopulated,water pollutionis River, andbaysclosetoKigomaonLakeTanganyika. Even downstream ofBrazzavilleandKinshasaontheCongo distinct impactsonrangerestricted species–forexample, downstream orlocalwaters.Insomeareas thesehavehad discharged directly backintoriversandlakes,polluting the alteration ofthemicrohabitats required by somefreshwater lakes, amajorthreat tomolluscsissiltation andsubsequent In tropical andcentralAfricainthe EastAfrica western 4.3.4 Sedimentation decline, increasing thealkalinityinlake. butitmayrapidlybecomeExtinctifwaterlevels Concern, occurs invastdensities,andiscurrently assessedas Least sp.”, isprobably alsoanendemic. to thislake,andasecondspecies,referred toas“ one. Onedescribedspecies, harbours agastropod fauna,albeitastrongly impoverished which isthemostalkalineofAfricanGreat Lakesthat still will decrease theamountofwaterreaching LakeTurkana, The plannedGibeIIIDamontheLowerOmoRiverinEthiopia is notalsothecaseinAfrica. (Pyron of thelocalextirpationgastropod populationsinthe USA particular reservoir assumed thathydrological alterationsfrom reservoirs, in in Africahavenotyetbeencarriedout,butitisgenerally of theeffects ofdamconstructionongastropod faunas appear tobetransportedbybirds (Van Damme1984).Studies Soapitia dageti Africa,suchas to western of thesespeciesare thoseingenerawhichare restricted regime andincreased watertemperature. Themostnotable mining wasteandsedimentation,alongwithchangingfl Côte d’Ivoire andGuineaare impactedbyacombinationof Some restricted rangeendemicspeciesfrom SierraLeone, in riversisdecliningduetotheexpansionofbauxitemining. are, however, quitevariable. problem throughout Africa.Thecausesofhabitatdecline Decline inhabitatqualityfreshwater riversandlakesisa 4.3.3 Water pollution species (Graf ofwaterfl patterns impacts onthehighdiversityhotspots,asanychangesto dams (seesection1.2,thisvolume)mayhavesignifi In centralAfrica,plansfortheGrandIngaandIII . Inmostcases,domesticwastewaterandsewageis et al. 2008),andthere isnoreason tosuspectthatthis andsome et al. (CR)( 2011).

ow andwaterqualitymayimpactthese release, are oneofthetwomaincauses García Gabbiella Sierraia outambensis

In western Africa,waterquality In western et al. Gabbiella rosea and Gabbiella rosea 2010;Kristiansen Hydrobia , is endemic , isendemic species that species that Melanoides Melanoides (CR) and (CR)and presently Tomichia et al. cant cant ow

and in Africa,andafurther three, Physa acuta Four species, Eleven ofthesespeciesoriginated inSouthAmericaorAsia. to fi reported inAfricawithin thepasttwodecades,belonging impacts. Fourteenaliengastropod specieshavebeen signifi Invasive molluscspeciesdonot,atthemoment,pose 4.3.7 impact onthemolluscfaunaisyettobeevaluated. aquaculture source asanalternative ofincome,butagainthe Africa hasledtochangesinlanduse,andtheexpansionof of galleryforest coveralongriversandstreams inwestern that haveconfi in thesehabitats,althoughthere havenotbeenanystudies Africa, haspossiblyhadanimpactonthenativemolluscs Victoria, andthedevelopmentoffi Africa,thevariousfi lakesofeastern tarn The stockingofnon-nativefi 4.3.6 Invasivespecies–fi snails ( also ledtoanincrease intheabundanceofbilharzia-carrying comm.) notedthattheincrease ininvasiveweedspecieshas snails ( are knowntoincrease habitatavailabletobilharzia-carrying molluscan faunaingeneralhasnotbeenevaluated,they Africa.Althoughtheirinfl southern present majorenvironmental andeconomicproblems in weeds (e.g., Herbert (1998)notedthatalienandhighlyinvasiveaquatic 4.3.5 extreme fl dwelling prosobranchs thatdonottoleratethechange to prosobranch species,butitcausesproblems forthebottom growth, whichmaybenefi decreasing stream velocitieswillalsofavouraquaticplant sediments foultheirgills.Increased exposure tosunlightand fi forest watersare alsoimpactedasthere isanincrease in Prosobranchs adaptedtolifeonhard surfacesinswiftrunning quantities offi chemical characteristicsofthesedimentsthrough increased the rivers.Theseaccumulationschangephysicaland forest cover, nutrient-enrichedsoilsrapidly accumulatein Melanoides of thebottomsediment,ortheylivewithinsediment(e.g., are benthicorepibenthic,tracingfurrows through thesurface snails. MostAfricanprosobranch species(unlikepulmonates) lms ofmudandfi ve families,twoprosobranchs andthree pulmonates. Gyraulus chinensis cant problems across Africabut dohavelocalized indigenous species Invasive species–molluscsandtheirimpacton Invasive species–waterhyacinthandotherplants Bulinus Bulinus uctuations in diurnal dissolved oxygen levels. dissolvedoxygenlevels. uctuations indiurnal species).Whenriverbanksare cleared oftheir and ne organic mudsanddepletedoxygenlevels. rmed this. As mentioned above, the clearance rmed this.Asmentionedabove,theclearance spp.and spp.and Pistia stratiotes Thiara granifera, Pseudosuccinea columella, Thiara granifera,Pseudosuccinea columella, Aplexa marmorata ne siltsthatcoverthesestones,andthe , couldpotentially become so,asthey Biomphalaria t most pulmonates and some small t mostpulmonatesandsomesmall Biomphalaria sh farming sh, suchastrout intheupper Pomacea difusa, R.rubiginosa and sh farminginwestern have become invasive havebecomeinvasive uence onthenative spp.)inLakeVictoria. Eichhornia crassipes Eichhornia spp.). Lange (pers. spp.).Lange(pers. sh farmedinLake )

CHAPTER 4 | MOLLUSCS 115

T. et 2010) of is present et al. 2009). Although probably has theprobably T. granifera T. et al. is often regarded as is often regarded at some sites. Another the native forms are being the native forms are 2004). The characteristic pattern on the indigenous gastropods of on the indigenous gastropods over pulmonates and may be the Tarebia granifera, Tarebia cant impact. In some cases, such M. tuberculatus et al. Tarebia granifera Tarebia falls within the wide range of characters has displaced the indigenous form in Lake has spread rapidly through the provinces of the provinces rapidly through has spread T. granifera T. T. granifera T. 2000), making it unlikely that its introduction . unpublished data). Where . unpublished data). Where et al has allowed it to invade several estuaries, lagoons et al. Melanoides tuberculatus, ow into southern seems no reasonow Mozambique. There M. tuberculatus M. tuberculatus 2010). Although these are species that should be monitored. It may be that It may monitored. should be that species are these Africa long been in have not invasives or all of these some serious as the extirpations have had impacts as enough to world, such as the Caribbean in other parts of the reported done to be made to research has recourse islands. Inevitably, it is unlikely but even here invasives elsewhere, on these same such islands small tropical in, for example, that observations to can be extrapolated directly as Puerto Rico and Martinique coming is, however, quantitative information Africa. Valuable ongoing study (Miranda N.A.F Miranda’s from the impact of the coastal lakes of northern Africa. KwaZulu-Natal, South major changes and species have caused Some introduced known aren’t others whereas impact on native species, to have had any signifi as For example,displaced by the south-east Asian morph. the south-east Asian morph has invaded Lake Malawi where southernit has displaced the indigenous form in the parts of the lake (Genner invasive morphof shell ornamentation of this sculpture and of 1994; (Brown seen in the polymorphic indigenous form Samadi will be easily noticed. Just as the invasive morphand spread of is evidence that it may also have happened inMalawi, there South Africa,western province, Africa and in KwaZulu-Natal years.in recent thiarid species Another parthenogenetic south-east Asian in the family Thiaridae, in large numbers, it is likely to have an important impact in large consuming as it does a substantialon primary production, which may explain the portion of the available food resource, dominance of cause of the loss of it is currently also moving southwards towards the Eastern towards moving southwards also it is currently it will and northwards, is predominantly Cape, its spread Mozambique and Zimbabwe soon if itundoubtedly reach northernmost so. In fact, the three rivers done has not already in which it has been found (Pongolo, Sand and Crocodile) all fl into eastern northwards why it should not continue spreading and central Africa. The high salinity tolerance exhibited by granifera and coastal lake systems in KwaZulu-Natal (Miranda al. quantities of benthic includes large its diet a detritivore, 68% of daily primary benthic productivity – up to microalgae (Miranda greatest potential of any invasive species to threaten native potential of any invasive species to threaten greatest Africa in 1999,species. Following its discovery in South granifera Tarebia as Swaziland (deKwaZulu-Natal and Mpumalanga, as well 2008; Appleton Kock and Wolmarans . Marisa Melanoides , have been Bulinus spp. spp. and and , has been recorded , has been recorded rmed by DNA analysis. 2010a). Pomacea uminea et al. Tarebia granifera Tarebia Corbicula fl Biomphalaria spp. , may represent an invasive morph of the species , may represent , and the thiarid ed as potential biocontrol agents of schistosomiasis ed as potential biocontrol © DANIEL GRAF AND KEVIN CUMMINGS Danger crocodiles! Field survey to be conducted with has care! are invasives in other countries. The other thiarid, are tuberculatus Asia, but this needs to be confi from seem as introduced Several species that have been reported bivalve, thenot to have survived. Only one alien freshwater Asian clam introduced Damme in African waters (Van cornuarietis identifi on or compete with host snails because they prey pulmonates such as Most invasive freshwater snails in Africa originated in tropical Most invasive freshwater of the world and, in terms of their regions or sub-tropical warm water species. Introductions thermal tolerances, are often (i) accidentally, can also be divided into those introduced (ii) deliberately, via the aquarium trade, and those introduced of the intermediate hosts ofusually for the biological control and, on transfer water schemes schistosomiasis. Inter-basin may also a much smaller scale, tanker transport of water, molluscs to new water bodies, as has distribute freshwater happened in Namibia (Appleton pers. comm.). Africa, widely across Some invasives have been recorded only a few or even singleknown from while others are countries. Despite the wide distributions of several species, quantitative information on their impacts on indigenous and often anecdotal. SeveralAfrican molluscs is scarce species, notably the apple-snails, Whether this competition is for resources, such as food, or Whether this competition is for resources, (e.g., physical contact) facilitated by thedue to interference Each case densities, is not clear. population invasives’ greater unique. Nevertheless, and more-or-less seems to be different 116 CHAPTER 4 | MOLLUSCS Children washingclothes inasteamneartheSanaga Children River inCameroon, potentiallyexposedtodiseases hosted by freshwater molluscs. hosted by freshwater © KEVINSMITH CHAPTER 4 | MOLLUSCS 117

, and cult 2005). , a (cause in Egypt, infections culties in were also were (cause of et al. Bulinus Biomphalaria S. intercalatum infections. © CHRIS APPLETON S. mansoni Physa acuta . S. haematobium , or both, in sub-Saharan Lanistes carinatus S. mansoni Schistsoma haematobium Calicophoron microbothrium Calicophoron S. mansoni uke as its snail host or (LC), a widespread species in southern (Gastropoda: Planorbidae), respectively, as Planorbidae), respectively, (Gastropoda: as well as the invasive as well as the invasive Bulinus is found the intermediate host species for urinary Bulinus Bulinus tropicus Africa and beyond is known to act as the intermediate host for the conical fl central nervous system and cause eosinophilic meningitis. cause eosinophilic system and nervous central reports parasite, isolated an Asian predominantly Although northern, westernfrom and southernAfrica suggest that in Africa become widespread may have angiostrongyliasis (1978) and Ibrahim (1975) and Yousif and Lämmler too. Yousif natural infections in reported species of experimentally that and showed and susceptible to infection. Schistosomiasis occurs in every country in Africa,Human schistosomiasis been known since the early decades except Lesotho. It has that the life-cycles of the two mostof the 20th century important schistosomes, schistosomiasis) and of urogenital involve species of intestinal schistosomiasis), Biomphalaria human parasite, intermediate hosts. A third people in parts of western and also uses affects Africa, species of infected with either The number of people, mostly children, S. haematobium 170 million, with another 600 Africa is estimated at around million at risk of infection (WHO 2002; Southgate estimated to die each 300,000 people are Approximately the disease (Hotez and Fenwick 2009), mostly year from in the case of kidney failure from and haematosis in the case of is diffi The economic impact of human schistosomiasis of diffi to gauge in monetary terms because of subtle variables like accurately measuring the effects educability and reduced absenteeism, loss of productivity costs Certainly, on a continent-wide or even national scale. of millions of US dollars per year. to Africa run into hundreds A widely used alternative of morbidity is the DALY measure common parasite of domestic animals. Within the genus Bulinus schistosomiasis, a serious human disease. are are

ukes. 2009). et al. Iridinidae)

. However, pea- . However, spp.). This factor . Rats are the usual . Rats are Neritina , but when people ingest ) are common in shallow common ) are T. granifera T. is sediment disturbance, as as disturbance, is sediment (1981) recovered 38 types (1981) recovered is worrying, since it will surely surely is worrying, since it will et al. rm their status as full species. cant in the future, as invasions by cant in the future, Pisidium spp. and T. granifera T. A. cantonensis and T. granifera T. Septaria Angiostrongylus cantonensis Sphaerium Understanding the impact of molluscs on human health and livelihoods nitive hosts for At present few freshwater molluscs are known to be directly known to be directly molluscs are few freshwater At present by climate change. In most cases, an increased threatened to events will be a secondary threat of drought frequency problems, already are there species where these freshwater of water for domestic and such as over-abstraction leading to low water levels in agricultural purposes, that are the rivers and springs. In western of water levels in Lake Chad Africa, the decline 1970s, leadinghas been consistent and ongoing since the in the size of the lake. This has led to the to the reduction category, listing of four endemic taxa within a threatened all in need of are although, as mentioned above, these taxa to confi taxonomic review 4.3.8 Climate change thiarids are documented to have resulted in great reductions reductions in great documented to have resulted thiarids are some cases, the extirpation of several in in numbers or, several Caribbean and bivalves in indigenous gastropods (Pointier 1999; Karatayev islands and the USA may become more signifi may become more disruption caused by caused disruption the sediment through its way that ploughs active snail it is an so. The deep as it does several millimetres leaving “trails” Unionidae and mussels (families freshwater Porter (1938) reported 112 morphologically distinguishable Porter (1938) reported across in 28 mollusc species from types of larval trematode South Africa, while Loker The economic importance of African freshwater molluscs The economic importance of African freshwater as intermediate hosts for parasites roles primarily on their rests causing disease in people and domestic stock. They serve fl as intermediate hosts for a wide variety of trematode of Tanzania. the Mwanza region in 14 mollusc species from cause economically important Several of these trematodes diseases in people and domestic stock, with those causing schistosomiasis and fascioliasis by far the most important. serve as intermediate hosts for the snails Many freshwater nematode 4.4 defi its larvae, either by eating infected snails or in mucus trails left by infected snails on salad material, they penetrate the unlikely to be severely affected, since they occur mostly in since they occur affected, be severely unlikely to by than is favoured water deeper endanger the prosobranch community there (including the there community endanger the prosobranch species of rarer clams ( exposed to impact. The invasion ofwater and could be along the south-east and eastthe brackish water zone coast of Africa by 118 CHAPTER 4 | MOLLUSCS the country, mostinthe NileDelta,with prevalences up An estimated830,000 peopleare thought tobeinfectedin Egypt where fascioliasisisregarded asanemerging disease. Human fascioliasis inAfricaneedsinvestigation. F. hepatica columella Studies elsewhere haveshownthattheNorthAmerican Africa, and gigantica liver fl Zambia. specimens afterafi Preserving death. years losttoadiseaseduemorbidity, disabilityandearly (disability-adjusted lifeyear),whichmeasures thenumberof Species of Fascioliasis more susceptiblethancattle. morbidity andmortalityhavebeenrecorded. Sheepare areas. Localizedoutbreaks accompanied bysignifi in death,withprevalence incattleofupto90%some S. margrebowiei distributed veterinaryspecies, as intermediatehosts,dothree otherlesswidely Both veterinary schistosomesare widespread overthecontinent. mattheei. mostly totheungulateschistosomes Veterinary schistosomiasis isalsocommon,anddue (Hotez andFenwick2009). malaria ortuberculosis, andalmostequivalenttoHIV/AIDS in Africaare ashigh70million–more thanlossesdueto ukes thatcausefascioliasis; S. bovis

Recent estimatesofDALYs losttoschistosomiasis Fasciola © DANIELGRAFANDKEVINCUMMINGS , the most common fasciolid in sub-Saharan , themostcommonfasciolidinsub-Saharan , whichisinvasiveinAfrica,susceptible toboth and Like the common human parasites, these Like thecommonhumanparasites,these L. truncatula Lymnaea F. gigantica infectionsare uncommon inAfrica,except and (Brown 1994).Heavyinfectionsmayresult S. mattheei serve as intermediate hosts for the serve asintermediatehostsforthe for ,

so itsrole inthetransmission of F. hepatica atLakeBangweulu, eld survey S. curassoni usespeciesof L. natalensis (Mas-Coma 2004). (Mas-Coma 2004). S. bovis , S. leiperi for and Fasciola Fasciola Bulinus Bulinus cant cant and and S. S. L. L. 4.5 diffi fascioliasis-endemic areas. Infectionsinsheepare more signifi effects are, however, diffi condemned liversandunderweightcarcasses. These rates. Themarketabilityofsuchanimalsispoor, dueto reduced growth, lowermilkproduction andlowercalving reported incattleandchronically infectedanimalsshow of stockanimals.Prevalences above50%are frequently and isknownacross thecontinentwhere itaffects arange Animal fascioliasisismore commonthanthehumandisease, bladder becomeinfl problem inEgypt.Inchronic infectionsthebileductandgall resulted inthediseasebeingrecognized asapublichealth the immature fl to 17%(Soliman2008).Thesevere pathologycaused by the value oftheirmolluscan biodiversity, socapacity relevant orindigenouscommunities appreciate governments importance ofmolluscs needstoberaised.Rarely dothe Capacity andawareness ofthevalueandecological are toprovide effective protection tofreshwater species. entire riverandlakesystems withintheirboundaries,ifthey specifi altered fl such aspollution,invasivespecies,sedimentation,and protected area boundarywillnotstopthe spread ofthreats pers. comm.),and,incaseswhere theyare included,the targeted conservationfeatures intheirownright(Darwall, rivers andlakesasboundarymarkersratherthan freshwater fauna.Themajorityofprotected areas include protected areas designedspecifi Africa there are, withtheexceptionofRamsar Sites,few protected areas asconservationactionsfor thespecies,in and thespring-snails,havebenefi world freshwater molluscs,suchastheriverrapidsspecies (Kristiansen andBrown 1999).Whilstinotherpartsofthe schemes toimprove thehealthoflocalpopulations eradication programmes aspartofbilharziamanagement conservation attention,more oftenbeingthefocusof of Africa.Infact,thefreshwater molluscfaunasgetlittle place toprotect anyofthethreatened freshwater molluscs There are noknowntargeted conservationmeasures in still applyonceadditionaldatahavebeenincluded. omissions, theconclusionsandsuggestedactionswould the threats tothesespeciesacross Africa.Evenwiththese representative ofthestatusmolluscanbiodiversityand described, webelievethatthedatapresented are reasonably and underestimates incurred aswewaitforspeciestobe At present, despitethefewomissionsfrom ourspecieslist cult toassess. measures Recommended conservation cant economiclossesare experiencedbyfarmersin cally toprotect uppercatchmentsandtoinclude ow regimes. Protected areas needtobedesigned ukes astheymigratethrough theliverhas amed aswell. cult toquantify. Nevertheless, ted from theextensionof cally forprotection of CHAPTER 4 | MOLLUSCS 119

as has et al. is usually Ferrissia, owing rivers cient to make cient to make eld-survey and and sh for the parasitic Burnupia (Iridinidae) is known Tarebia granifera Tarebia ed from South America ed from . Most of these species Burnupia cation services, as well as cient species cient cient, awaiting the results of cient, awaiting the results . (2011) note, in western Africa Ancylus cations are currently best made currently cations are c impacts of dam construction, c impacts Mutela joubini et al eld, which may explain why they are why they are eld, which may explain ve museum lots, mostly in the Chad-Chari has recently been identiﬁ been has recently shing or aquaculture on these bivalves (Graf shing or aquaculture Research studies on species demography and demography studies on species Research ecology Field survey for Data Deﬁ

v) Control of invasive species of invasive v) Control to better understand the impacts of invasive Research where species, especially in regions become established, should be a priority. larvae of freshwater mussels of the families Unionidae, mussels of the families Unionidae, larvae of freshwater minimal, prohibiting are Iridinidae and Etheriidae evaluation of the specifi overfi mussels could also native freshwater 2011). The decline in changes in the river ecosystems, as lead to fundamental water purifi the mussels provide mussels (e.g., habitat for commensal species living on the acari). Many such studies could be suitable as freshwater if small sources student projects and MSc. undergraduate of funding could be provided. iv) known from are molluscs in the region Many freshwater small number of specimens, and most of only a relatively collecting. by recent those have not been corroborated For example, as Graf mussel the freshwater only fi from Basin of the western As another example, African region. by 49 species the family Ancylidae is represented belonging to two main genera, well as a few species of assessed as Data Defi are As these freshwater by C. Albrecht. ongoing research likely to be overlooked by very small they are limpets are collectors in the fi so poorly known. Identifi to the type localities of the on the basis of proximity the genus described species. Finally, to be indigenous to Africa, but a species of considered Burnupia by dos Santos (2003), raising biogeographical questions as to the true distribution of the genus. Species such in need of further fi clearly as these are to make informed conservation in order taxonomic review poorly and management decisions, especially in the more of Africa, such as the smaller lakes in surveyed regions eastern Africa (e.g., Lake Albert), the fast-fl in western and parts of the Congo basin where Africa, described in the many of the least known species were last century. that any proposed developments in these regions take regions in these developments proposed that any species. endemic of these account iii) taxa in Africa, demographic, molluscan For many freshwater insuffi are and life history data ecological bivalve and about how gastropod meaningful predictions environments. to changing freshwater species will respond on the host-fi For example, information

sh et al. 2008), et al. lling the niche vacatedlling the niche eld survey or taxonomic ow for conservation sheries departments, are recommended recommended are sheries departments, sh. c community has led to pressure to publish c community has led to pressure Decline of food supplies for the other native species Decline of food supplies for the other native on these molluscs, such as crabs, aquatic that predate and fi birds by the non-carrier species; and Increase in the carrier species, fi Increase Facilitate information fl management

are less diverse than some continental faunas, the current less diverse than some continental faunas, the current are some are There number of species is underestimated. and eastern such as the north African Maghreb regions, species many recognised are there Africa lakes, where that have yet to be named (Ghamizi 1998; West in western also regions and central Africa are 2003). There has been little active fi there where under- will be since the 1960s, and these regions research and may potentially support unrecognised recorded estimates of species diversity may species. The current The over the next 10 years. need revision well, therefore, lack of “value” placed on new species descriptions by the scientifi overview papers or synthesis papers, rather than traditional This, combined with a decline in journals alpha-taxonomy. papers dealing with descriptions of new that will take large species, is undoubtedly slowing the speed with which new described. species are ii) that information on range-restricted It is critical to ensure authorities, and endemic species is passed to the relevant Although the freshwater molluscs of Africa, with an Although the freshwater estimated 560 species (Bogan 2008; Strong i) Description of new species farm developments; large-scale timber extraction involving timber extraction farm developments; large-scale and mining developments of gallery forests; clear-felling methods. using open surface extraction of molluscides needs to be betterThe indiscriminate use also causes decline of populations as it currently controlled, a threat species that do not provide of non-carrier endemic of these species to human health or livelihoods. The loss has two major effects: a) building projects, such as is ongoing in Morocco (IUCN in Morocco as is ongoing such projects, building withcollaboration and outreach community through 2010) and fi local wildlife of local monitoring and facilitate to raise awareness bivalves. and of gastropods populations include need to Impact Assessments Environmental and impacts to mollusc diversity, assessments on the developments likely for any proposed should be mandatory such as: dam construction; fi to impact mollusc species, Finally, sewage treatment needs to be improved, and there and there improved, needs to be sewage treatment Finally, on the import of invasive species needs to be tighter control the impact on native species. to reduce actions 4.5.1 Recommended research b) 120 CHAPTER 4 | SPECIES IN THE SPOTLIGHT @ P.O. Box150, Curepipe,Mauritius ² School ofBiological andConservation Sciences,University ofKwaZulu-Natal, Durban,4000 SouthAfrica ¹ Photo 1.TheAfrican openbillstorkwithitscharacteristic “open”bill. \jg\Z`Xccpkfk_\8]i`ZXefg\eY`ccjkfib =i\j_nXk\idfccljZjXi\`dgfikXekÆ stork, they aretotheAfricanopenbill we canseeclearlyhowimportant be conserved,butinthisshortessay molluscs areimportantandshould and mussels.Inparticular, itfeeds exclusively onfreshwatersnails The openbillstorkfeedsalmost some caseevenwithinit)that conservation community(orin convince peopleoutsideofthe t isnotaneasytaskto Anastomus lamelligerus Jg\Z`\jèk_\jgfkc`^_k . family Ampullariidae( on largeoperculatesnailsofthe as well asunionoidanbivalvessuch on and does abirdusegreatbigbeakto also taken.Thequestionis:how as theAchatinidae,aresometimes (Iridinidae). Terrestrial snails,such wahlbergi (Unionidae), and Unio Bellamya Pila

caffer spp.),andlesscommonly and spp.(Viviparidae), as and Mutela zambesiensis Chambardia Coelatura framesi © DAVID ALLAN Appleton, C.C. ¹andLaHaussedeLalouvière,P² Lanistes ovum

shells werefound onfl Delta, accumulationsof near Chief’sIslandintheOkavango this unresolvedquestion. bills, presentsnewhypotheses on shells withtheshapeofbirds’ comparison ofthedamageto KwaZulu-Natal, SouthAfrica,and fl Delta, Botswana,andthePongolo openbill storksintheOkavango Examination ofshellsdiscardedby on somekindofsolidsurface. always withthepreyresting amongst vegetation,butapparently it isusuallydoneunderwateror and diffi to removethesoftpartsisintricate The processofmanipulatingshells prior toremovingtheirsoftparts. manipulate individualmolluscs like pincersandpickup the tipsofmandiblestooperate shells, butisproposedtoenable thought tobeusedforcrushing for SouthAfricanstorks,isnot the mandibles,5mmonaverage The characteristicgapbetween from theshellsoftheirprey. the birdstoextractsoftparts is theactualtechniqueusedby the subjectofdebate,butsoalso of thegapinstorks’billstill 2006). Not onlyisthe function 1960; Kahl1971;Hockey their molluscanprey(e.g.,Huxley in extractingthesoftpartsfrom that theiconic“open” billplays feeding behaviourandtherole reviewed literatureonthestork’s papers onAfricanbirdshave shells? mussels frominsidetheirprotective extract thebodiesofsnailsand oodplain innorth-eastern On twooccasionsinJuly2000, Various booksandscientifi cult toobservebecause attened L. ovum et al c .

CHAPTER 4 platforms, approximately 40cm across, made by openbill storks at a height of 30-40cm above the water, in clumps of the reed Vossia

cuspidata (Photo 2). A total of 46 |

shells were collected from these SPECIES IN THE SPOTLIGHT platforms, and all were large adults of 40 to 55mm shell height. These shell larders seem analogous to the accumulations of empty shells reported from river banks in eastern Africa, and suggest that the openbill stork reuses the platforms as surfaces for extracting the soft parts from prey snails over a period of time – a sort of kitchen workbench! No bivalves were Photo 3. L. ovum collected from the reed platforms in the Okavango Delta, Botswana, found here, but since they live in showing the triangular notches in the lip of the basal whorl. © CHRIS C. APPLETON deep water, they are inaccessible to the birds. Curiously though, no Pila occidentalis, a second large storks’ lower mandible which is pushes the sharp lower mandible ampullariid occurring in the delta also triangular in cross section. through the notch to sever the (Appleton et al. 2003), were present A comparison of these notches operculum and then the columellar even though they grow to 55mm with the bills of museum specimens muscle, probably in one movement. and form part of the openbill’s diet of the openbill stork suggests The lower mandible of the openbill elsewhere in Africa. that the birds use a sophisticated stork is, in fact, slightly curved, Most shells from these platforms technique to extract the soft parts and Kahl (1971) proposed that this (72%) had a single more-or-less of snails like L. ovum. Having curvature made it easier for the triangular notch in the apertural picked the shell up, and secured bird to sever the snails’ operculum lip 5 to 21mm (mean 15mm) deep it between the tips of its grooved and columellar muscle. Five of the (Photo 3). A further six had shallow upper mandible and the knife-like museum specimens examined had chipping and the remaining seven lower mandible, the stork uses the the lower mandible curved to the showed only slight irregularities to lower mandible to take a triangular right, one to the left, and three the outer lip – not very different notch out of the outer apertural lip were not curved at all. Nevertheless, from empty shells found elsewhere to expose the operculum. Using the this action frees the snail’s soft in the delta. Allowing for uneven upper mandible to hold the snail parts from its shell so that they can breakage of the shell, the shape and fi rmly on a surface like the reed be eaten by the bird. depth of these notches match the platform described above, it then A similar technique is used by openbill storks for bivalves or mussels, but with the difference that bivalve prey is initially held crosswise in the bird’s bill. This allows the stork to chip into the ventral margins of both valves with its lower mandible. It then fl icks the shell into a lengthwise position with its dorsal hinge line lodged in the grooved under surface of the upper mandible (or puts the mussel with its notched valves onto the bank), inserts its lower mandible between the valves via the notch and, starting at the posterior end and drawing it towards the anterior end, severs the two adductor Photo 2. An openbill storks’ larder – a platform in a clump of Vossia cuspidata with muscles as it does so. Once these numerous empty Lanistes ovum shells. © CHRIS C. APPLETON muscles have been cut, the soft 121 122 CHAPTER 4 | SPECIES IN THE SPOTLIGHT concluded thattheopenbill stork Huxley (1960), had previously numbers consumedinthiscase. quantitative datatoestimatethe openbill storks,butwehaveno on thefl unpubl. data). in 1983(P. laHaussedeLalouvière, counted onthefl fl fl openbill storksonthePongolo a substantialpartofthediet suggest thatbivalvesrepresent 591,560 individuals).Thesedata standing cropof individuals) and4.6%ofthe crop of consumed 13.0%ofthestanding We estimatethatopenbillstorks caffer Unio Floodplain inKwaZulu-Natal. Ch. wahlbergi standing cropof predation byopenbillsonthe we haveassessedtheextentof la HaussedeLalouvière(1987), for 1983/4fromAppletonand markers, andbivalvedensitydata to bivalvepreydescribedaboveas (Photos 4candd). taken outoftheposteriorend of bothspecieshadshallowchips 4a-d). Asmallernumberofvalves on thePongolo fl framesi valves ofapproximately60% the ventralmarginsofarticulated present neartheposteriorendsof as deepin shells areremarkablyclean. inner surfacethatthediscarded close totheirscarsontheshell’s These adductormusclesarecutso parts canberemovedandeaten. Near Threatened in SouthAfricatheendemic ocks comprisingalmost450birds oodplain, withanestimated20 Various authors,including Using thecharacteristicdamage Triangular notches,seldom and oodplain, andiseatenby Co. framesi in declineworldwide,and Unionoidan bivalvesare Ch. wahlbergi L. ovum onthePongolo L. ovum oodplain’s lakes oodplain (Photos Ch. wahlbergi Co. framesi (or27x106 , werealso alsooccurs collected isclassifi and (or Co. (Appleton 1985). framesi (probably synonymouswith shells of people inZambiatoopensimilar the techniqueusedbyIronAge hit withabluntobject,whichwas crushing. Crushingisakintobeing different fromthatresulting damage describedherearequite to doso,butthetypesofshell the gapbetweenitsmandibles crushed theshellsofitspreyusing the valves. umbonal areas and anteriorendsof by fragmentsincorporating the bivalves wererepresentedmostly their aperturesintact,whilethe parts oftheirbasalwhorlbutkept had typicallylosttheirspiresand from thesearchaeologicaldeposits shallow chipsatthevalves’posteriorends. notches intheposteriorhalfofvalves’ventralmargins,whilecanddshow (b) valvesfromthePongoloRiverfl Photo 4(a-d).Examplesofnotched c (d) (b) (c) (a) ed as ) and L. ovum Ch. wahlbergi , Co. mossambicensis L. ovum shells , forfood Co. oodplain. Photosaandbshowcharacteristic Co. framesi © CHRISC.APPLETON snail golf coursesforthecommongarden foraging inbuilt-upareasandon range insouthernAfrica,and openbill storksexpandingtheir in birdwatchers’blogsitesof This mayexplainrecentreports mussels’ parasiticlarvalstage. species thatserveashostsforthe invasive fi molluscides, andtheimpactof increasing pollution,useof Threatened, probablydueto Unio caffer and inSouthAfricatheendemic bivalves areindeclineworldwide, openbill stork?Unionoidan dependent speciessuchasthe such asthesepotentiallyimpact important species. the fateofthese unobtrusivebut decisions which mightdetermine understanding tothosewhomake The challengenowistospread this apparent, excepttoaselectfew. in wayswhichmaynotatfi is importantafterall,andoften conclusion, thehumblemollusc indigenous achatinidstoo.In So howdothreatstomolluscs (a,candd) Helix aspersa sh ontheindigenous isclassifi Ch.wahlbergi andprobably ed asNear rst be CHAPTER 4 Jg\Z`\jèk_\jgfkc`^_k :Xek_\cXjkgfglcXk`fejf]k_\^`Xek |

8]i`ZXe]i\j_nXk\ig\Xicdljj\cY\jXm\[6 SPECIES IN THE SPOTLIGHT

Ghamizi, M¹ and Van Damme, D²

or decades this large mussel associated to another host, and it was believed to be Extinct, is feared that the same thing will until surveys in the area happen to M. marocana, which may rediscovered it just a few result in it being lost forever. =years ago. Formerly, M. marocana All habitats in Moroccan rivers was considered to be a local race from which populations of M. of the European Margaritifera marocana have been recorded during auricularia, found over a large the early 20th century are heavily part of western Europe. However, degraded, threatened by desiccation a few specialists, such as Dr. Margaritifera marocana (CR) is a large and polluted by domestic and Douglas Smith, disagreed with naiad species endemic to north-west agricultural effl uents. Some this view. But on the basis of shell Morocco. Moroccan types of Margaritifera, characteristics from a handful of such as the one originally described museum specimens collected at as M. dernaica from the Oued Derna, the beginning of the 20th century, gills of their host fi sh, and after are completely gone. Are M. dernaica this taxonomic problem could metamorphosing into a young and M. marocana one and the same not be solved. It was only after mussel, detach and drop to the species? Only in the unlikely event live specimens were rediscovered sediment. This way, the fi sh ensures that live Margaritifera specimens three years ago that DNA research the distribution of the clam. Host would be rediscovered in the Oued by Araujo et al. (2009) provided specifi city in all Margaritifera Derna could this question be solved. the irrefutable evidence for species is high, restricted to a single The conservation of M. marocana, Margaritifera marocana’s specifi c or a few fi sh species. It is not yet considering the recent IUCN distinctiveness. The species is, known which and how many fi sh assessment of its threatened status, hence, a Moroccan endemic, species are used by M. marocana as warrant investigations concerning whose range is restricted to two hosts. If its host specifi city is very not only the biological questions rivers belonging to the same narrow, it may be that the particular already cited above, but should hydrographical basin. The area that fi sh host has disappeared from the also include sociocultural research they occupy within these rivers is rivers where they are known (for on the Amazigh communities that extremely limited. So far, only two example, as a consequence of the reside along the river at the two sites small populations have been found, construction of dams), therefore where Margaritifera still occurs. both at sites strongly affected by preventing reproduction and the Nowadays, local children swim, human activities. Both populations completion of the life cycle of particularly during summer, at consist exclusively of specimens of M. marocana. Virtual complete both sites without paying attention old age-classes, all shells measuring extinction of its main host fi sh, to the mussels, which look like more than 10cm. With no evidence the Atlantic sturgeon, led to the fl attened stones, wearing sandals of juvenile mussels, recruitment extirpation of the European to protect their feet against cuts appears to have been halted, and Margaritifera auricularia except in from the valves’ edges. In 2009, recruitment patterns remain an two rivers where populations are when collecting material for unknown. The same is true for the species So far, only two small of fi sh host or hosts on which the larvae of M. marocana are parasitic. populations have been As with other representatives of the genus Margaritifera, the found, both at sites strongly larvae (of the glochidium type) affected by human activities attach themselves to the skin and

¹ Faculté des Sciences, Marrakech, Morocco ² Faculty of Sciences, Ghent University, Belgium 123 124 CHAPTER 4 | SPECIES IN THE SPOTLIGHT continued existence. we shouldallbe concernedforits species isCritically Endangeredand the problemofrecruitment.This species inordertofi our biologicalresearchonthis also equallyimportanttocontinue conservation ofthespecies,but itis campaigns aresetupforthe imperative thatawareness swept awayorburiedinsediment. fl desiccation interruptedbymassive that causesperiodsofdroughtand precipitation andriverfl the increasingirregularityof and climatechanges,inparticular by thedegradationofitshabitat disappearance beinghastened presently dyinginsilence,its should discoveranyuseforthem. last remainingpopulationsifthey with nohesitation,exploitthe two that thevillagerslivingnear zone. Yet itshouldbe keptinmind to theinhabitantsofcoastal Morocco seemstobeconfi oysters andotherseaproducein two sites.Consumptionofclams, the villagesinvicinityof freshwater mussels,atleastnotin appears tobelinkedthese as well,noculinarytradition tourist carsandbuses.Fortunately is alsoamainstoppingpointfor important siteforthesemussels especially giventhatthemost of pearl),forsaletotourists, with theirbeautifulnacre(mother shells (uptoalengthof20cm), offering theselargeriverpearl relief toseethatnoonewas several occasions,itwasagreat the mussels. the potentialcommercialvalueof youngsters werewonderingabout consisted ofEuropeans,thelocal to thefactthatpartofteam about theirinterestbecause,due mussels. Thereweremixedfeelings team andhelpedbydivingfor enthusiastically joinedtheresearch genetic research,thesechildren oods, during whichmusselsare It is,therefore,notonly Margaritifera marocana On returningtothesesiteson Margaritifera siteswould, nd solutionsfor ow regime is ned : inkling oftheirtruedistributions that theyexistandhaveonlyan of species,wearemerelyaware mussels ingeneral.For themajority is typicaloftropicalfreshwater know about is stillagreatdealthatwedon’t the dryseason.However, there to survivethepunishmentof attributed totheanimal’sability mollusc-hostile continent)canbe success ofthismollusc(inatruly the continent.Thegeographical recorded frommostregionsacross to particularbasins, have smalldistributions,restricted mussel speciesintheAfrotropics Africa. Whilst themajorityof study the evolutionary relationships of African freshwater mussels. ofAfricanfreshwater relationships study theevolutionary Y`kf]dljZc\ 8dfccljZn`k_X University of Alabama ¹ Chambardia wahlbergi, Jg\Z`\jèk_\jgfkc`^_k species ofmolluscin of themostwidespread a freshwatermussel,isone hambardia wahlbergi C. wahlbergi C. wahlbergi collectedfromafi , andthis (LC), is sheries pond. This specimen will be used to sheries pond.Thisspecimenwillbeusedto the Africanfreshwatermussel belongs. Relativetootherregions, family that only fromAfrica,anditistothis Iridinidae. Theiridinidsareknown Margaritiferidae, Etheriidae,and families occurinAfrica:Unionidae, in sixfamilies.Four ofthese more than850speciesworldwide dinosaurs, andtodaythereare originated atthesametimeas group ofbivalvemolluscs.They globally distributed,andancient Unionoida belongtoaspecies-rich, environment (includingus). these molluscsinteractwiththeir inhibits ourunderstandingofhow living musselsinAfrica,andthis detailed fi been collected.Itisraretohave based ontheplacestheyhave Freshwater musselsoftheorder eld observationsfrom Chambardia wahlbergi © DANIELL.GRAF Graf, D.L. ¹

CHAPTER 4 |

SPECIES IN THE SPOTLIGHT

The Lutemwe River (Luangwa Basin) in Zambia at low water. © ALEC LINDSAY fauna is restricted, with only about upon freshwater fi shes. It is very 80 species (compare this to 300 in likely that C. wahlbergi larvae do the North America, or more than 200 same. In 2008 and 2009, with the in south-eastern Asia). Africa is help of Alex Chilala of the Zambian a tough place for molluscs, with Department of Fisheries, it was waters poor in calcium (needed for possible to collect C. wahlbergi from their shells), and the dry climate artifi cial ponds stocked with tilapia. limiting for aquatic organisms. This provides strong evidence that Chambardia wahlbergi ranges the larval mussels had hitchhiked from South Africa north to along with the parental stock Somalia and west to Senegal. In of fi sh, but the dynamics of the southern and eastern Africa, C. reproductive cycle of this mollusc wahlbergi is able to thrive where remain to be explained. other mussels cannot because of To the uninformed, freshwater its ability to survive when rivers mussels like Chambardia wahlbergi dry up seasonally. In 2005, samples may be mere “clams”, but they were taken from the Lutemwe have the potential to capture the River of Zambia. During that time Chambardia wahlbergi, collected from imagination of anyone open to of year, the stream is reduced to a a fi sheries pond. © DANIEL L. GRAF acknowledging the complexity mere trickle, but with the generous and interrelatedness of aquatic assistance of the local people who ecosystems. However, these animals use the subsurface waters for both Unfortunately, most of our are threatened worldwide due to irrigation and laundry, several knowledge of the biology of the degradation of fresh waters. living specimens of C. wahlbergi Chambardia wahlbergi is limited to Basic biological research on the were obtained from deep within such anecdotes. We can only infer freshwater molluscs of Africa will the sediment. With at least 40% its life history from better-studied be crucial for establishing informed of Africa dominated by desert- freshwater mussels, assuming conservation priorities, not only for like habitats, this ability gives C. a great deal from the traits of the protection of biodiversity but wahlbergi, and freshwater mussels temperate species. For example, for the sustainable management of like it, a distinct advantage. almost all mussel larvae are parasitic fresh waters into the future. 125 Acisoma panorpoides (LCRG) a widespread species found Chapter 5. across Africa, and southern Asia. © J.-P. BOUDOT Dragonfl ies and damselfl ies of Africa (Odonata): history, diversity, distribution, and conservation

Dijkstra, K-D.B.¹, Boudot, J-P.², Clausnitzer, V.³, Kipping, J.4, Kisakye, J.J.5, Ogbogu, S.S.6, Samraoui, B.7, Samways, M.J.8, Schütte, K.9, Simaika, J.P.8, Suhling, F.10 and Tchibozo, S.L.11

1 Netherlands Centre for Biodiversity Naturalis, P.O. Box 9517, NL-2300RA Leiden, The Netherlands 2 LIMOS - UMR 7137, Universités de Nancy, Faculté des Sciences, Boulevard des Aiguillettes, BP 70239, F-54506 Vandoeuvre-lès-Nancy Cedex, France 3 Senckenberg Museum of Natural History Görlitz, PF 300154, D-02806 Görlitz, Germany 4 Naturkundemuseum Mauritianum Altenburg, Parkstrasse 1, D-04600 Altenburg, Germany 5 Department of Zoology, Makerere University, P.O. Box 7062, Kampala, Uganda 6 Aquatic Entomology, Department of Zoology, Obafemi Awolowo University, Ile-Ife 220005, Osun State, Nigeria 7 Laboratoire de Recherche et de Conservation des Zones Humides, Biology Department, University of Guelma, Guelma, Algeria 8 Department of Conservation Ecology and Entomology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa 9 Biozentrum Grindel und Zoologisches Museum Hamburg, Entomology, Animal Ecology and Conservation, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany 10 Institut für Geoökologie, Technische Universität Braunschweig, Langer Kamp 19c, D-38106 Braunschweig, Germany 11 Centre de Recherche pour la Gestion de la Biodiversité et du Terroir, 04 B.p. 0385 Cotonou, Benin CONTENTS 5.1 Introduction 128 5.1.1 Odonata in biogeography and conservation 128 5.1.2 Knowledge of African Odonata 128 5.1.2.1 Taxonomy and identifi cation 128 5.1.2.2 Distribution and ecology 129 5.2 Diversity and history 132 5.2.1 Diversity 132 5.2.1.1 Richness 132 5.2.1.2 Patterns of diversity 132 5.2.2 History 133 5.2.2.1 Relevance and context 133 5.2.2.2 Impoverishment and diversifi cation 133 5.2.2.3 Relict fauna 138 5.2.2.4 Modern fauna 141 5.3 Regional diversity, endemism and threat 143 5.3.1 Afrotropical fauna 143 5.3.2 Guineo-Congolian fauna 145 5.3.2.1 Overview 145 Notogomphus cottarellii 5.3.2.2 Upper Guinea 147 (EN) is an Ethiopian endemic 5.3.2.3 Lower Guinea 148 threatened by habitat 5.3.2.4 Congo Basin 149 destruction. © K.-D.B. DIJKSTRA 5.3.3 Afro-montane fauna 153 5.3.3.1 Overview 153 5.3.3.2 South Africa 154 5.3.3.3 Angola 156 5.3.3.4 Eastern Mountains and Coast 156 5.3.3.5 Ethiopia 158 5.3.4 Complex faunas 159 5.3.4.1 Albertine Rift 159 5.3.4.2 Zambezia 160 5.3.5 Insular fauna 162 5.3.5.1 Indian Ocean 162 5.3.5.2 Atlantic Ocean 164 5.3.6 Palaearctic fauna 164 The globe skimmer 5.4 Conservation 166 (Pantala fl avescens) (LC). 5.4.1 Status 166 © VIOLA CLAUSNITZER 5.4.1.1 Red List assessment of African dragonfl ies 166 5.4.1.2 Distribution of threatened species 167 5.4.2 Major threats 168 5.4.2.1 Habitat degradation and deforestation 168 5.4.2.2 Dams 169 5.4.2.3 Mining 169 5.4.2.4 Alien plants and fi sh 170 5.4.2.5 Water extraction and pollution 170 5.4.3 Research actions 170 5.4.3.1 Field survey 170 5.4.3.2 Ecological research 171 5.4.3.3 Understanding change 172 5.4.3.4 Information disclosure 172 5.4.4 Conservation actions 173 Species in the Spotlight – A guardian of Kenya’s watershed: Maathai’s longleg 174 Species in the Spotlight – The globe skimmer: on a journey around the world 175 Species in the Spotlight – The Pemba featherleg: a fragile damselfl y in a fragile realm 176 Species in the Spotlight – A wonderful water relic Hadrothemis vrijdaghi with an ancient lineage 177 (LC). © K.-D.B. DIJKSTRA 128 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA from collections andthefi a thirdspeciesin the complex,(c) genetically. of onespeciesuntilMitchelland Samways (2005)demonstratedthey differed asforms treated they were Insharpco ingenitalmorphology(Karsch1894),sizeandwingmarkings (O’NeillandPaulson2001),pruinositypatt Despite differences (a) 2007d) anddistributionsare suffi other Afrotropical insects(seereviews inDijkstra2003b, Odonate taxonomyiswell-resolved incomparisonto 5.1.2.1 Taxonomy andidentifi 5.1.2 KnowledgeofAfricanOdonata and Suhling (2005), Dijkstra conservation andincorporatingdatafrom Darwall as anelaborationofDijkstra(2007d),focusingalsoon rapidly changingcontinent.Anoverviewisprovided here, Odonata willhelpusunderstandthepastandfuture ofa human, geologicalorclimaticinfl changing nature oftheAfricanlandscape,beitunder tropics, andcertainlyinAfrica.Consideringtheever- sources, isaproblem world-wide,especiallyinthe catchments, withtheconsequentlossofsoilandwater- vital tothesurvivaloflife.Humandisturbancewater habitats where waterisscarce and,therefore, especially habitats suchaswetlandsandrainforests, butalso for not onlyasfl 2008). Duetotheirattractiveappearance,theyfunction Jödicke 2004;Clausnitzer scientists onregional andgloballevels(Clausnitzer and increasing attentionfrom thepublic,conservationistsand Oertli 2008;Samways2010;Suhling Clausnitzer 2003;Corbet1999;DijkstraandLempert (conservation), aboveandbelowthewatersurface(e.g., long term(biogeography, climatology)andtheshortterm diversity. Theyare suitedforevaluatingchangeinthe a vitalhumanresource andthelargest class of animal conspicuous freshwater insects,thusrepresenting both Dragonfl 5.1.1 Odonatainbiogeographyandconservation 5.1 Introduction a b (c) (b) (a) Palpopleura lucia is n damselfl and ies et al. agships forconservationofwater-rich et al. (2009a). (LC)and(b) (2010;2011),Samraoui Biodiversity Assessment. eld during andafterthe IUCNFreshwater ies (Odonata)are themost et al. P. portia P. albifrons cation 2009;Córdoba-Aguilar uence, thestudyofits (LC) were thefi (LC)were et al. ciently knownfor (LC),wasonlydescribed fromGabonin1979.Thefi 2006),receiving et al. rst odonatestobenamedfromAfricansoil,in1773(Dijkstra (2010) et al.

dragonﬂ obscured thestatusofsomemostfamiliar family, Libellulidae,where prolonged taxonomicinertia progress, isillustratedbyAfrica’s mostconspicuous The needfortaxonomicadvancement,andits (2) (1) problems havebeencentraltothetaxonomicdisorder: be published(DijkstraandClausnitzerinprep.). Two males ofovertwo-thirds ofthecontinent’s specieswill stride forward shouldbemadein2011,whenakey to which hasimproved surveyingintheregion. Afurther and Martens2007;Tarboton and Tarboton 2002,2005), partofthecontinent(Samways2009;Suhling southern Several keysandﬁ problems orthelackofregional data(see5.1.2.2). our understandingisstillpartlyobscured bytaxonomic biogeographic studiesandthreat assessments,although

by fragmentarycollecting. continent withfewnaturalbarriers,andisexacerbated geographic andenvironmental rangesacross ahuge great inAfrica,asmanyspeciescanrealise broad forms andsubspecies.Theproblem isespecially but hasalsoledtothedescriptionofsuperfl age-related. Thisappliesmainlyatthespecieslevel, is probably environmentally induced,butcanalsobe 2003a, 2005a,Dijkstra but are notsupportedbymorphology(Dijkstra Many supposedtaxadiffer insizeandmelanism, ecological adaptation. of veinsinducedbychangesinwingshapeasan as primitivemayratherbederivedreconfi believed. Charactersthatwere traditionallytreated quantifi Vick 2006).Venation wasfavoured becauseitis by othercharacters,suchasgenitalia(Dijkstraand of more taxa(especiallygenera)thanare supported An overvaluationofwingvenationledtotherecognition ies,moststrikingbeingthe‘former’ able,butismuchmore plasticthanpreviously eld guidesare already availableforthe ©

W. TARBOTON (a,b) ANDN.MÉZIÈRE(c) rst records since were only obtained onlyobtained since were rst records et al. 2006a).Suchvariation et al. Palpopleura 2003). 2003). gurations ern, ern, ntrast, ntrast, uous . CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 129 (south). (south). 2010). 2010). B. leucosticta et al rst continent-wide, high-resolution, ed database of a group of tropical of tropical ed database of a group (north) and (north) demonstrate that even inspection of “common” and demonstrate that even inspection of “common” and species is warranted (Dijkstra and “recognisable” Matushkina 2009; Damm 5.1.2.2 Distribution and ecology the Odonata Database Developed as part of this project, of Africa (ODA) is the fi taxonomically-verifi Brachythemis impartita Brachythemis remained remained is one of is one of B. impartita were only discovered once that once that only discovered were Brachythemis leucosticta Also, while most familiar odonates, Africa’s unnoticed among it for 170 years, despite having a 5.1; Dijkstra and widely overlapping range (Figure large “within” and Matushkina 2009). Structural differences stictica Trithemis “species” was found to be genetically heterogeneous (Damm and Hadrys 2009). These and further cases Adapted from: Dijkstra and Matushkina (2009). and Matushkina Dijkstra from: Adapted Figure 5.1. Inferred distributions and the overlap for overlap and the distributions 5.1. Inferred Figure 130 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA Figure 5.2.Densityoflocalityrecords ina100x100kmgrid.Adaptedfrom: Kipping Democratic Republicof Congo(D.R.C.), Mozambique research incountrieslike Angola, RepublicofCongo, were studied relatively constantly through time,whereas and space(Figures Africa 5.2 and5.3).Partsofsouthern resolution isalready impressive, dataare patchyintime It nowcontainsover75,000 records, butwhilethe ofdiversityandresearchdetermine patterns intensity. freshwater insects, thus providing auniquetoolto 5.2.2.1; 5.4.3.2-3). providing thefi rst ecological dataonthespecies(see place, improving distributionalknowledge andoften historic records, asurge inrecent fi eldwork hastaken but havenotyetbeendatabased. Asidefrom databasing gaps). Datafrom some regions, likeNigeria,are available independence (see5.4.3.1for anoverviewofrecording and Somaliawasimpeded by politicalunrest after et al. (2009). CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 131 eld survey survey eld of fi the intensity in ecting variation ect the efforts of Elliot C.G. Pinhey, and the 1980 to 2010 records to 2010 records and the 1980 C.G. Pinhey, of Elliot efforts ect the (2009). et al. (c) (d) (a) (b) over time. 1920 to 1949 is a period of high collecting activity during colonial times, especially in especially times, colonial during activity of high collecting a period to 1949 is time. 1920 over largely refl to 1979 records D. R. C.; 1950 during that time. and political unrest subject to civil war for those areas lack of information highlight a Kipping Adapted from: Figure 5.3. Density of locality records over four time periods refl time periods over four locality records of 5.3. Density Figure 132 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA Table 1.Comparisonoftropical odonatediversity. Datafrom Kalkman Neotropics, butare represented byonlytwospecies Perilestidae andPseudostigmatidaeare shared withthe With bothregions itshares allbuttwoofitsfamilies; only 16families,compared to21forbothotherregions. many species.TheAfrotropical faunaisclassifi both theNeotropical andOrientalregions havetwiceas total represents lessthanone-sixthoftheworldfauna; and aboutanother50inPalaearctic northAfrica. The the surrounding islands(mainlyMadagascanendemics) must lieabove725species,withabout150more on undescribed species,thetotalintropical mainland impoverished atalltaxonomiclevels(Table 1).Including The Afrotropical odonatefaunaiscomparatively 5.2.1.1 Richness 5.2.1 Diversity 5.2 Diversityandhistory aiygnr pce eeaseisgnr species genera species genera species genera Family oa 8 661286251665 235 816 112 1636 186 Total eauia – – 1 1 Petaluridae eptlia – – 1 1 Neopetaliidae armia 6250 2 36 1 2 2 Macromiidae iellde4 5 6275 190 56 217 46 352 44 Libellulidae opia 6231 3 3358 43 134 15 273 26 Gomphidae poheide––11 1 – – Epiophlebiidae odlia 761 57 7 19 6 37 2 Corduliidae odlgsrde11–527 5 – 1 1 Cordulegastridae hooopia 40 1 – – Chlorogomphidae utoeaide27–– – 7 2 Austropetaliidae ehia 51753 8138 18 39 5 127 15 Aeshnidae ANISOPTERA ylsia 7 1 2 9 2 1 1 Synlestidae suotgaia 811– 1 1 18 5 Pseudostigmatidae suoetde––315 3 – – Pseudolestidae rtnuia 49 3857 8 23 2 94 14 Protoneuridae oyhrde85 – – 58 8 Polythoridae ltsitde14 119 5 – 42 1 Platystictidae ltceiia 58130 8 45 9 – Platycnemididae eietde21 – 1 1 18 2 Perilestidae eaoaroia 41053 0298 10 38 5 130 14 Megapodagrionidae etiede––14 1 – – Lestoideidae etde23 4539 5 14 1 38 2 Lestidae uhede––1 65 12 – – Euphaeidae itraia – – 2 2 Dicteriadidae onginde3 7 1192 185 23 179 11 370 38 Coenagrionidae hooyhde–34 480 14 43 3 – Chlorocyphidae aotrgde36 71 60 10 17 3 61 3 Calopterygidae mhpeyia 5 1 2 1 3 2 Amphipterygidae ZYGOPTERA etoisArtoisOriental Afrotropics Neotropics e into ed and KwaZulu-NatalontheIndianOcean(Figure 5.4a extends from GuineaandAngolaontheAtlantictoKenya as averyrich(more than100specieslocally)area that in thenorth(Sahara)andsouth(Namib-Kalahari),aswell reveals twovastareas withfewerthan25specieslocally The distributionofodonatediversityintropical Africa ofdiversity 5.2.1.2 Patterns American andAsiantropics. the largest odonatefamiliesonEarth,isaparwiththe Afrotropical diversityofCoenagrionidaeandLibellulidae, of thegreater similaritybetweenAfricaandAsia.Only the groups throughout thePalaeotropics isfurtherevidence et al. Old andNewWorld “protoneurids” are notrelated (Carle Moreover, macromiids donotoccursouthofMexicoand Platycnemididae shared withAsiarepresent almost11%. (0.2% ofthefauna),whileChlorocyphidae and 2008),andtherefore therichrepresentation ofthese et al. (2008) andDijkstra(2007d). CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 133 2008), Africa’s 2008), Africa’s cation et al. 2006, 2009b, 2010; 2006, 2009b, 2010; ies to arid and seasonal ies to arid and seasonal fth centre of diversity and of diversity and fth centre et al. 2010; Suhling et al. with an important centre of endemism, the Upper Guinea, of endemism, the Upper Guinea, with an important centre respectively, Lower Guinea, Albertine Rift and Zambezia, combined with highlands leads to high rainforest where and thus high species diversity (see habitat heterogeneity 5.4b). A possible fi 5.3.2; Figure Basin appears endemism lacks highlands; the Congo but 125 species locally, comparatively poor with around of Africa (see it includes some of the least surveyed parts 5.3.2.4). 5.2.2 History 5.2.2.1 Relevance and context ecological amplitudes, and thus their Species’ current determined to complex impacts, were response future 2010). An by their past (e.g., Samways and Niba aids the conservation understanding of history therefore to future by anticipating response diversity of present change. For example, long distance dispersal evolved in to deforestation enables a response to aridity response of too. Africa can be seen as the “experimental garden” climatic pronounced global change, experiencing more drier towards with a trend shifts than other tropics, see Dijkstra 2007d). The conditions (for references, of dragonfl ecological response become better understood (e.g., conditions has recently Martens certain Huyssteen and Samways 2009). For example, Van in dry years, expanding to wetter areas species retreat wet years (Samways 2010). again in outwards 5.2.2.2 Impoverishment and diversifi drier Extinction caused by unstable and progressively impoverishment (see climates explains much of Africa’s harbours most of the rainforest 5.2.1.1). As tropical odonate diversity (Kalkman world’s and so its impoverishment reasonable is quite proportion

cult cult cant cant Urothemis in press), and and in press), gure suggests a suggests a gure and (b) et al. fths of the national fauna is Pseudagrion © N. MÉZIÈRE (a) (b) rst, then drops sharply where the tropical African fauna African fauna the tropical sharply where rst, then drops from Gabon. overleaf), with rather abrupt diversity gradients between between overleaf), with rather abrupt diversity gradients hyper-diverse them. The gradients north and south of the although that is diffi belt seem to be shaped differently, of data. In to demonstrate with the available resolution desert ends (more the north, diversity rises steeply where begins (more forest than 25 species) and again where diverse fauna than 100), with a rather uniform and equally In the south, in the savannah and woodlands in between. the extend far from rivers, swamps and gallery forest at diversity diminishes gradually southwards wet centre; fi highlands. meets the Kalahari desert and South African the Okavango This is demonstrated in Namibia, where 100 species, the savannah have around and Caprivi areas highlands 71 woodlands of the dry central Namibian deserts species, and the hyperarid Namib and Kaokoveld (Suhling only 23 and 11, respectively two-fi in South Africa, where north-east. This pattern limited to the tropical is masked parts of which Angola, large by the lack of data from In the east than 100 species locally. may have more the diversity pattern is complex; the fi the western the ocean, but while rift towards from drop truly probably are high and dry parts of Tanzania large and Mozambique is impoverished, much of Tanzania simply unstudied. coincides with the continuous Although the richest area of western lowland rainforest and central Africa, the eastern a diversity and stability of highlands provide habitats that allows it to extend well east and south. local diversity where areas, four core the Moreover, than 200 species, each include signifi more reaches The species richness of the families (a) Coenagrionidae and (b)The species richness of in Africa. This diversity includes theseLibellulidae is unparalleled of (a) striking unnamed species highland areas: (1) west, from Sierra Leone to Togo; (2) (2) Sierra Leone to Togo; west, from (1) highland areas: to Republic of Congo; (3) north-east, Nigeria central, from including Uganda and north-east D.R.C.; and (4) south- overlaps east, including Katanga and Zambia. Each core 134 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA present aridity, itsmissingfamilies maycome from its While Africa’s lower speciescountisrelated topastand below), forexample,canbefound incloseproximity. local diversity:one-third ofall odonate specieshavelarge ranges on average,buthigh African tropics (Fjeldså 2003;Plana2004).Also,African lowland rainforest have localdiversitycomparabletonon- endemism, large partsoftheAfricansavannahand overall impoverishmentresults from less narrow geographically more uniformfauna.AlthoughAfrica’s ranges andmore widespread competition, leadingtoa may havegreater dispersalcapacities,enablinglarger Moreover, inresponse to(recurrent) aridity, Africanbiota can beconsidered asmuchcontemporaryhistoric. Odonata. Figure of(a)speciesrichnessand(b)rangerestricted 5.4.Patterns species(ranges<50,000km²)ofAfrican (a) Trithemis species(see moister centre andwest.In thecaseof Africa,ratherthaninthe the speciesoccur ineastern America andAustralasia (Corbet1999).Notably, twoof are known(Figure 5.5overleaf),muchlessthanin notably rare. Forexample, onlyfourtree-hole breeders to desiccation(e.g.,seepages, temporarystreams) are climatic history. Specialistsofforest habitatsvulnerable Africa’s lossoffamiliescanalso beexplainedbyits retreating intoAfricaduringcolderperiods. Nonetheless, isolation impededEurasianwarmtemperatetaxafrom many biotathatradiatedatthetime(Morley2000).This were tropical andincontact,thus“missed out”on 54 millionyearsagowhenmostotherlandmasses until theclosure oftheTethys Sea.Itwasisolated60- relative isolationsincethebreak-up ofGondwanaland Coryphagrion , CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 135 Hadrothemis Hadrothemis © K.-D.B. DIJKSTRA (LC). H. coacta (LC) this adaptation may have evolved from a species reproducing in forest pools. The latter habitat is evidenced by the (a) (b) (b) Breeding in tree-holescomparatively scarce among African Odonata: in the golden-winged Congo basin endemic (a) is muddy abdomen tip of this female (b) vrijdaghi 136 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA fi Figure 5.5.Distributionofdendrolimnetic Afrotropical Odonata.Legend–opencircles: From: Dijkstra(2007d). extensive extinction,it alsogeared therise ofspecies if speciationrates were lowerand changeinduced 2000; Plana2004; White2001).Nonetheless, even habitat complexity(Fjeldsåand Lovett1997;Morley varied relief (e.g.,fewer mountainchains)andassociated the break-up ofGondwanaland andtherefore hasless landmasses, Africahasbeen geologically stablesince from alowerspeciation potential;compared toother extinction andlowerratesofcolonisation,butalso A lowerspeciescountcanresult from higher ratesof but were subsequentlyextirpated. (2009) suggestedthatplatystictidsoriginatedinAfrica, and Petaluridaehavespecialisedecologies:vanTol Megapodagrionidae, Platystictidae,Pseudostigmatidae four otherspecies.Thescarce orabsentfamilies bred intemporaryforest pools,thehabitat ofthegenus’s the three dendrolimnetic stable environments (see5.2.2.3),buttheancestorsof may beproof ofsurvivalinonethecontinent’s most coconut husks(ClausnitzerandLindeboom2002),this which evenbreeds inwaterheldsnailshellsand lled circles: Hadrothemis scabrifrons Hadrothemis speciesprobably ; halvedcircles: bothspecies;shading: et al.

diversiﬁ and Clausnitzer(2006)Dijkstra(2007d)formulated as newhabitatsbecameconstantlyavailable.Dijkstra that straddledenvironmental anddispersalbarriers giving rise tomanylineages inlittletime. Asopenhabitats forest expansionseparatedpopulations inopenareas, period between 5and3.5millionyearsago. Possibly et al. is bestadaptedtoliveindesert environments (Suhling remainder of conditions. Indeed, favoured thoseadapted toexposedandtemporary while dryingdisadvantagedmanyaquaticspecies,it stagnant habitats,coincidingwithsavannahexpansion; inferred anorigin6to9millionyearsagoinopen and geographicaldata(Figure 5.6),Damm molecular phylogeny(81%ofspecies)withecological and from lowlandstohighlands.Combiningadated to warmtemporarypools,from deserttorainforest, with aboutfortyspecies,from coolpermanentstreams genus dominatesdragonﬂ and Trithemis, 2009b).Themainradiationcoincided withawetter cation scenariosfor Trithemis butonlythelatterhasbeentested.The H. camarensis T. kirbyi (excluding theatypical y communitiesacross Africa , thesister-species ofthe Coryphagrion grandis Coryphagrion Platycypha and H. vrijdaghi , Pseudagrion et al. T. brydeni (2010) . ; ),

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 137 T. (a)

2010). et al. 2008). 2008). y radiation: et al. adult habitat: water; (b) standing owing vs. is exemplary of an African dragonﬂ open to forested habitats and from standing to running standing to running habitats and from open to forested at least eight and six times, respectively, waters occurred and two times. For example, only three the reverse expansion 1 million years ago perhaps provided forest for two species access to suitable shaded streams now found in lowlands. that are of the highland group diversity in Africa appears to be dominated Present-day with such high adaptation potentials. The by groups coenagrionid and libellulid abundance, being the families not surprising. Only best adapted to change, is therefore by the ice ages, impacted strongly in temperate regions, is their dominance and the impoverishment of all other families comparable to that in changeable Africa (Dijkstra and Clausnitzer 2006; Kalkman Trithemis T. ies, ies, © C. VANAPPELGHEM (a), K.-D.B. DIJKSTRA (b) AND E.R. NIELSEN (c) © C. VANAPPELGHEM and (LC). T. annulata T. T. tropicana T. (c)

(LC) and ed excessively in the next 3.5 million T. aenea T. ), (b) RG ve more dispersals to Eurasia or Madagascar dispersals to Eurasia or Madagascar ve more , two of Africa’s most widespread dragonfl widespread most , two of Africa’s (LC (a) (b) (c) ow, these lineages barely radiated further. By contrast, By contrast, further. radiated these lineages barely ow, which are currently expanding northwards in Eurasia. in Eurasia. expanding northwards currently which are At least fi as a demonstrating the variable environment occurred, of history cradle for adaptable species and the actuality with such potential for gene in modern change. However, fl within the ecological constraints of lineages three and swampy habitats highland streams, lowland streams, diversifi respectively, Shifts from 55% of species diversity. years, producing arteriosa began to coalesce after 3.5 million years ago, the species began to coalesce after 3.5 million years into largely adapted to temporary conditions expanded overlapping ranges. These include arteriosa The diversity of ecologies and colorations exhibited by the genus some forest cover vs. open habitats. Based on Bayesian 50% majority-rule consensus phylogram obtained from from obtained phylogram consensus majority-rule 50% on Bayesian Based open habitats. cover vs. forest some from posterior probabilities node indicate the Pie charts on each and ITS regions. data set of ND1, 16S combined see Damm (for details using SIMMAP state reconstructions analyses of ancestral the stochastic Figure 5.6. Ancestral state reconstructions for (a) larval habitat: fl larval habitat: for (a) state reconstructions 5.6. Ancestral Figure 138 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA Pseudostigmatidae; S:Synlestidae). area isindicated(A:Amphipterygidae;M:Megapodagrionidae; Pe:Perilestidae;Pl:Platycnemididae;Ps: Figure 5.7.Distributionofrelict Afrotropical Zygoptera.Thenumberofspeciesandthefamilyforeachgenus/ Paracnemis Corduliidae to thesister-group ofallLibelluloidea(= Macromiidae, (Figure 5.7),aswell three genera(probably) belonging undetermined relations nowplacedinPlatycnemididae small (mostlymonotypic)restricted-range generaof Perilestidae, PseudostigmatidaeandSynlestidae,several members ofAmphipterygidae,Megapodagrionidae, spray (Vick 1998).Thepresumed relicts includeAfrican Stenocnemis lack closerelatives withinthecontinent.Anexampleis relatively smallorisolatedranges,andthat(largely) should focusonecologicallysensitivetaxa,whichhave probable centres ofclimaticstability. Thecomparison the oldestAfrotropical species,theirpresence indicating faunas ofMadagascarandothertropical areas mayidentify A comparisonofthecontinentalAfricanOdonatawith 5.2.2.3 Relictfauna Phyllogomphinae, represented by on thecontinent, suchastheendemicgomphid subfamily are taxa well-represented bothinMadagascarand locally 5.8). Strong candidates tobequalifi , andLibellulidae;seeWare (Dijkstraunpubl.).Adaptedfrom: DijkstraandClausnitzer (2006). , whichinhabitsrock faceswettedbywaterfall ed as“oldAfricans” Isomma Nesolestes nigeriensis et al. 2007)(Figure (including Malgassogomphus Neotropics. Dumont represented bytwogenera and18speciesinthenorthern of theGuianaShield. Besides relative of Australian (Dijkstra Synlestidae and of therelict Afrotropical Odonataare unclear. Thoseofthe (5) thegraniticSeychelles(5.3.5.1). Theaffi Arccoastal andEastern forestsAfrica(5.3.3.4); ofeastern and surrounding equatorialrainforest (5.3.2.3); (4)the towards theCape(5.3.3.2);(3)LowerGuineahighlands rainforests (see5.3.5.1);(2)SouthAfrica,especially importance: (1)Madagascar, especiallytheeastern be identifi Five centres where these“oldAfricans” survivedcan Madagascar(compareand eastern Figure 5.8). confi and thelibellulidgenera equatorial Africaand isincludedin ned toforest streamsAfrica incentraland western ed (Figures 5.7-5.8,5.10,5.13),inorder of Pentaphlebia Neurolestes Syncordulia ) onMadagascar, et al. et al. Ceratogomphus Malgassophlebia isthemonotypicgenus 2007c;Ware (2005)dated the , ofSouthAfricaare possibly Metacnemis secundaris Nubiolestes et al. in southern Africa, insouthern Phyllogomphus and 2009).Theonly , Perilestidaeis nities ofmost Pentaphlebia Neodythemis Rimanella in in - , CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 139 and ; star: both ; star: both Idomacromia Idomacromia I. proavita Nesocordulia (NA). 2007). 2007). S. legator et al. Ware ; standing triangle: ; standing sensu I. lieftincki (2007c). . ; inverted triangle: ; inverted et al (LC), and four Syncordulia species, like this (b) I. jillianae ). From: Dijkstra ). From: Chlorolestes umbratus Chlorolestes I. proavita species overlap too widely to be indicated separately (species number provided); number provided); (species separately to be indicated too widely overlap species and (a) (b) © W. TARBOTON (a) AND M. SAMWAYS (b) (a) AND M. SAMWAYS TARBOTON © W. South Africa is home to the greatest concentration of relict odonates in continental Africa. Among these are nine synlestids, represented here by (a) I. lieftincki I. lieftincki Figure 5.8. Distribution of (probable) Afrotropical “GSI-corduliids” ( “GSI-corduliids” Afrotropical of (probable) 5.8. Distribution Figure Syncordulia separated (diamond: species are 140 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA the twothinlinesreceives >250mmofrainduring bothhalvesoftheyear. From: Dijkstra(2007d). rainforest RiftValley. duringtheEocene,darkshading zonesofupliftintheEastern Thearea between Figure 5.9.Biogeographichypothesisfor Walvis Ridge, whichconnect southBrazil, through the America andAfrica wasalongtheRioGrande Riseand Coast forests. Apossibledispersal route betweenSouth habitats ofNeotropical pseudostigmatidsthantheEast rainforests bearagreater resemblance totheperhumid submontane forest streams, contemporaryLowerGuinean 1992b; ClausnitzerandLindeboom 2002),ratherthanin breed inlowlandforest phytotelmata(Fincke1992a, not surviveintheeastaswell.Althoughpseudostigmatids survived (seeabove)andraisesthequestionwhythesedid vicariance impliesextinctionwhere otherNeotropical relicts from themonotypic 5,500km ofAtlanticOceanand2,500kmAfricaninterior – 18Neotropical species,infi The biogeographyofpseudostigmatidsismore perplexing (cumulatively andperennially) wettestpart(Maley1989). in continentalAfrica,where theirrangecoincideswiththe these damselfl distribution, supportGondwananvicariance,highlighting Rimanella split around 150millionyearsago.This,andtheir ies aspossibly theoldestsurvivingodonates Coryphagrion ve genera,are separatedby (Figure 5.9).Gondwanan Coryphagrion . Thepaleshadingrepresents theapproximate extentof of Tanzania was littleinfl fragmentation) rainfall fl to short-(desiccation ofhabitats)andlong-term (forest the poordispersalofadults, (Lovett 1993).Giventhesmallvolume oflarvalhabitatsand barrier towestward dispersal by18-17millionyearsago (Morley 2000).Upliftandassociated aridifi forestssouthern disappeared around 36millionyearsago years ago,allowing forest extendedalongtheIndianOceanabout55million equatorial forests bymore openlandscapes. However, roughly oppositetheWalvis Ridge,were separatedfrom forests,the equatorialone(Morley2000).Southern circumglobal bands,withtwosubtropical bandsfl (Parrish 1993).Atthistime,rainforests extendedinthree opportunities forisland-hoppingpersistedwellthereafter may havebeenlostbefore theCretaceous, (diminishing) million yearsago.Althoughanintercontinental connection about 34millionyearsago,theRisesubsidedaround 25 (McDougall andDuncan1988).TheRidgesubmerged present-day hotspotatTristan daCunha,tonorthNamibia Coryphagrion uctuations. Thecoastalclimate uenced byPleistocene changes, Coryphagrion tomovenorthbefore the cation created a isvulnerable anking CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 141

is ius Ă Sapho nities. has a cation, . also took shared Prodasineura shared (34 myo) from (34 myo) from wings (about 25 wings (about and shared Arabineura khalidi Ictinogomphus y Neophya Umma Prodasineura and Ictinogomphus Ischnura Phaon iridipennis Arabicnemis caerulea Elattoneura and and and and cation (see 5.2.2.2). They are all are (see 5.2.2.2). They cation Gomphidia Sapho , ed in few genera) species, suggesting suggesting species, genera) ed in few Aciagrion Elattoneura Gomphidia ius and Nel 2009). Dijkstra (2007d) inferred (2007d) inferred ius and Nel 2009). Dijkstra Ă (2005) estimated that ancestors of (2005) estimated that separated from Asian stock 77-85 million Asian stock 77-85 separated from et al. Diastatomma ; ** Umma nities are alleged nities are million years old) from France and million years old) from England (Nel and Paicheler 1993; Nel and Petrulevi England (Nel and Paicheler 2010; Petrulevi exchange was associated with the that African-Eurasian ago, Sea 24-10 million years of the Tethys gradual closure Madagascar. virtual absence from conforming to their Dumont and but that is questionable given their place in the Cretaceous, The paradox of Miocene dispersal morphological similarity. aridifi into Africa is that it coincided with extensive of a marine especially after 16 million years ago: the closure Sea, lead to the opening of a terrestrial the Tethys barrier, specialists either the Sahara. African rainforest hurdle, ancestors of open or evolved locally from arrived earlier, habitats (see 5.2.2.2). The damselfl oases, but is closely related endemic to Arabian stream to the diverse complexes of species in Africa and Asia. years ago and that these genera split 49-53 million years years ago and that these African rainforests in been present ago, but had they These Madagascar. have reached they should this early, continental and authors believed that the separation of Madagascan populations of enigmatic affi similar range and habitat, but more of the African-Eurasian Both species could be relicts Odonata. stream exchange of tropical and Clausnitzer 2004). All examples have many relatively relatively have many examples 2004). All and Clausnitzer (i.e., classifi similar rapid and recent diversifi recent rapid and Asia, although faunas in tropical diverse to more related warmer Eurasia in across have ranged widely they would fossils showing Afrotropical times. European and wetter affi Proischnura

and et al. Species Madagascar, Species Madagascar, and MascarenesComoros Relatives in southern Asia Pinheyagrion , from western from Brazil Ischnura , , arriving from Africa and , arriving from 17 - none known 1717 - - none known but genus not shared many, Species African continent liation of the 120 million years liation Azuragrion Euarchistigma , Coryphagrion provides some test of its credibility. A credibility. some test of its provides Chlorocnemis 2006). A-group B-group 41 22 - endemic, 1 shared 1 closely related many in genus, probably not related many in genus, probably (2008) demonstrated that pseudostigmatids (2008) demonstrated Africallagma and

, et al. et al. Coryphagrion Aciagrion CalopterygidaeChlorocyphidaeischnurine Coenagrionidae*Pseudagrion 35Pseudagrion 17Allocnemis 43 genera in shared 5 endemics with continent 1 shared many, - genera but no shared many, genera but no shared many, Protoneuridae 23 - many, Notogomphus Macromiidae 35 genus 1 endemic in shared genera but no shared many, lindeniine Gomphidae**Neurogomphus 13 - many, unlike further south in Madagascar’s rain-shadow (Fjeldså (Fjeldså rain-shadow in Madagascar’s further south unlike genus’s 1993). The Lovett Lovett 1997; Fjeldså and 1994; precipitation perennial coincides with relatively range still frame falls within a time 5.9). The above scenario (Figure estimate of the divergence years and only an of 100 million of dated (Groeneveld phylogeny was not molecular 2007) and the family afﬁ 2007) and the family is doubted (Bechly pers. comm.). (Carle and Wighton 1990) Carle no but provided nested within the Coenagrionidae, are anomaly, time. As a biogeographic estimate of divergence the are and caviomorph rodents monkeys the New World image of mirror old “pseudostigmatid” in South America only 26-25 and beginning to radiate and Wyss 1998; (Flynn respectively 31.5 million years ago, Opazo 5.2.2.4 Modern fauna nowadays dominant in None of the “old Africans” are large even absent from continental Africa and they are the Guineo-Congolian Especially areas. hyper-diverse fauna is dominated by “new Africans” (see Dijkstra forest absent on Madagascar and Clausnitzer 2006), which are at most by a few adaptable colonists (Table or represented 35% of Afrotropical 5.2). The examples alone represent of which (i.e., including Madagascar) species diversity, examples 97% is continental. By contrast, the “old African” with only 48% 5.7-5.8 form 12% of diversity, in Figure continental continental species. The distinction between species, and insular faunas is less clear for open-land better dispersers, but most of those occurring which are arrivals (Dijkstra recent considered on Madagascar are * Table 5.2. Examples of large (>10 species), morphologically homogeneous groups, which are widespread in widespread which are Examples of large (>10 species), morphologically homogeneous groups, 5.2. Table Dijkstra (2007d). From: (largely) absent on Madagascar. often Asia, but are Africa and tropical (a)

Two monotypic genera endemic to stream oases in southern Arabia may represent relicts of odonate exchange between tropical Africa and Asia: (a) Arabineura khalidi (VU) and (b) Arabicnemis caerulea (VU). © B. REIMER

(b) CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 143 , Z. and by fth Tramea Tramea Acisoma , avifrons , Orthetrum Rhyothemis Hemistigma Agriocnemis , , avescens Chalcostephia fl Tholymis tillarga , Zygonyx natalensis , Urothemis edwardsii Urothemis edwardsii fths. The name of the , Pantala fl Ischnura senegalensis , , and Diplacodes lefebvrii , , ne affi

. Some mainland species are replaced replaced are . Some mainland species H. Aethriamanta rezia Tetrathemis polleni Tetrathemis , by , Trithemis arteriosa Trithemis Palpopleura lucia , , Ceriagrion glabrum , and generally low habitats, such as savannahs. Many of savannahs. such as habitats, low and generally throughout occur and species genera pan-African these the wettest,possible exception of with the the Afrotropics, Eurasia may extend well into and coldest areas, driest and pan-African with purely islands. Countries or to oceanic nations such as the Sahel very impoverished, faunas are each. The dominance fewer than 100 species with mostly fauna Libellulidae in the Afrotropical of Coenagrionidae and among 5.2.1.1; 5.2.2.2) is even greater (50% of species; see being fauna (70%), examples this widespread exilis trinacria panorpoides Crocothemis erythraea Zygonyx torridus such as in Madagascar by a close relative, albipunctum semihyalina basilaris fth cult. Diversity fth is associated fth of diversity is (Figure of richness regions ed three fths of species occur predominantly fths of species occur predominantly cation of both regions and species diffi cation of both regions Regional diversity, endemism endemism diversity, Regional threat and 5.3.1 Afrotropical fauna 5.3.1 Afrotropical often adaptable species, barriers, and With few geological making biogeographic is extensive, range overlap classifi 5.3 (see 5.2.1.2.), for which Dijkstra and in forests is greatest Clausnitzer (2006) identifi or highlands, but with open is not associated with forests 5.10). Almost two-fi of the western lowland forests andwithin the extensive 5.3.2). Just over a fi central continent (see with the eastern and southern part dominated by highlands meet ‘high Africa’, ‘low Africa’ slopes up to (5.3.3). Where a “mixed” character occur (5.3.4)complex faunas with 5.11-5.12). About another fi (Figures fi in Madagascar (5.3.5). The remaining centred insular, (about 150-175 species) of the total species richness, the thick-bordered ellipse two-fi richness, the thick-bordered (about 150-175 species) of the total species in given in lower case. Regions are by each ellipse is given in upper case; discussed regions fauna represented in italic font have a 5.7-5.8); those endemic genera (Figures often in regionally bold font harbour ancient relicts, Dijkstra (2007d). 5.11-5.12). From: faunas intersect (Figures “mixed” character where Figure 5.10. Schematic representation of Afrotropical odonate diversity. Each ellipse represents about one-fi Each ellipse represents odonate diversity. of Afrotropical 5.10. Schematic representation Figure 144 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA circles: knownsitesoftheAfro-montane a few species survive inisolated suitablepockets,like(b) a fewspeciessurvive TanzaniaSouth oftheSahara, openswampsasinwestern pan-Africanspecies. Inthenorthonly (a) canharbouralmostevery demarcates theAlbertineRift.Legend–shading: approximate rangeoftheGuineo-Congolian Figure 5.11.Distributionof AND B.SAMRAOUI (b) a (b) (a) Notiothemis species.Theoverlapofthesetwoshade-dwellingspeciesapproximately N. jonesi. From: Dijkstra(2007d). Urothemis edwardsii ( LC RG ) innortheastAlgeria. © K.-D.B.DIJKSTRA (a) N. robertsi ; CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 145 D. ve allopatric species, but in the north. Legend – open in the north.

complex can illustrate the distribution Allocnemis- Note the concentration of the pan-African the concentration Note has a gradual transition of riverine forests and woodland has a gradual transition of riverine forests into peripheral savannahs. The highest diversity in the 200 all countries with well over is found here: Afrotropics within species have a considerable portion of this forest and Clausnitzer 2006). Although (Dijkstra their borders it can region, the forested many species range throughout of endemism (Figure be subdivided into four main areas 5.13). The odonates because all species are and diversity of forest (Figure obligate dwellers of shaded permanent streams in the Guineo-Congolian is widespread 5.14). The group up to four species overlap and even up to where forest, highlands. The Afro-montane seven in the Cameroon by ﬁ is inhabited archipelago in the northernmost is absent the group sites, despite the habitats (see 5.3.3.5). Further detail of suitable presence of radiation Saphoinae, an Afrotropical by the is provided and streams to forested 15 calopterygid species restricted on the highlands in Diversity is centred 5.15). rivers (Figure – Upper Guinea, with the Lower and – to a lesser degree as only single species in the Guineo-Congolian periphery, . . Diplacodes. ; stars: both species. Adapted from: Dijkstra (2006a; 2007d). Dijkstra (2006a; from: species. Adapted ; stars: both , and Azuragrion Z. lachesis T. donaldsoni T. D. pumila with A. kauderni and Bradinopyga strachani with (2010; 2011), Samraoui , in the Lower Guinea and (2009a) for further details, et al. et al. Z. fuelleborni G. quarrei ; closed triangles: ; closed triangles: cient species and regions. cient species Trithemis dejouxi Trithemis and , which generally inhabits smaller and more elevated swamps, elevated and more inhabits smaller , which generally A. buchholzi and A. nigridorsum , and and in the Congo Basin (see 5.3.2.4; Figure in the Congo Basin (see 5.3.2.4; Figure (2005), Dijkstra . The northern and south-eastern savannah in the large swamps in the north and south of the Paleo-Chambeshi region (shading) and that of Afro- and (shading) region of the Paleo-Chambeshi south north and in the large swamps in the D. deminuta D. pumila

et al. (2010) and Suhling B. cornuta Zygonoides fraseri Both latter examples also have isolated siblings within – matrix the forest Z. occidentis and endemism and threatened 5.17). Regional diversity, See discussed below. (NT) species are Near Threatened Darwall and Such pairs with also a Madagascan relative are are Such pairs with also a Madagascan relative et al. especially of Data Deﬁ 5.3.2 Guineo-Congolian fauna 5.3.2.1 Overview Central and western dominated by almost Africa are which continuous Guineo-Congolian lowland rainforest, vansomereni elisabethae belt may also harbour two vicariants, for example, Gomphidia bredoi Figure 5.12. Distribution of small continental African African continental of small Distribution 5.12. Figure deminuta montane triangles: 146 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA C. maccleeryi maximal numberofsympatricspeciesindicated);squares: shading: rangeof Figure 5.14.Distributionof thickness ofborder indicatesrelative importance(Figures 5.14-5.15).From: Dijkstra(2007d). bordered ellipses:approximate positionofAfro-montane centres ofGuineo-Congolianodonaterelicts, black-bordered ellipses:approximate positionofcentres ofancientrelict Odonata(Figures 5.7-5.8);grey- dark shading:approximate positionofmainPleistocene rainforest refuges, astraditionallyidentiﬁ distribution ofGuineo-Congolianforest Odonata, dividedintofourmainareas ofregional endemism; Figure 5.13.Afrotropical forests andtheirpresumed refuges. Legend –paleshading:approximate current ; triangles: Allocnemis leucosticta C. marshalli Allocnemis . Adaptedfrom: DijkstraandClausnitzer(2006).

and ; paleshading:rangeofGuineo-Congolianspecies(regional Chlorocnemis (including C. abbotti; Isomecocnemis circles:

C. montana ). Legend–dark ; diamond: ed;

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 147 (a) Allocnemis (LC) are © J. KIPPING (a) Chlorocnemis witteiChlorocnemis LC) and (b) AND E.R. NIELSEN (b) (a) ( leucosticta two southern African representatives of a platycnemidine radiation concentrated in the Guineo-Congolian forests. The accents of bright colour on their bodies stand out in their gloomy habitats. of number indicate the gures (b)

C. , and and 2010). combined. Shading intensity and ﬁ intensity Shading combined.

(VU) and Elattoneura Elattoneura et al. U. declivium , Diastatomma in Angola and Phyllogomphus Umma ava , (NT) are known only known only (NT) are M. fl and and Trithemis africana Trithemis avipennis , Agriocnemis angustirami C. fl Chlorocypha luminosa Platycnemis guttifera in Angola and , , , Sapho Elattoneura dorsalis are present throughout in forest in forest throughout present are (CR) from at most a few sites in at most (CR) from . The threatened and NT endemics and . The threatened (EN), U. femina Mesocnemis dupuyi Sapho ciliata , with a sympatric species pair in western in Tanzania (Dijkstra and Vick 2006). (Dijkstra and Vick in Tanzania Lestinogomphus matilei Phyllomacromia sophia , , Prodasineura villiersi (CR) and Chlorocnemis elongata , Mesocnemis tisi , well as the endemic and central Africa, and the endemic in the Eastern highlands. A similar pattern is shown by Arc Micromacromia M. miraculosa 5.3.2.2 Upper Guinea occur in Nigeria and further west, than 300 species More endemic (Dijkstra 60 are than of which more Many such as radix Phyllogomphus moundi of and Lempert 2003) and therefore (Dijkstra streams Least Concern. The same applies for strict Upper Guinea such as endemics, not occurring east of Togo, gamblesi aethiops Zygonyx chrysobaphes localised and rather dispersed: more are pluotae watershed, and the Gambia from girardi (VU), Pseudagrion mascagnii Figure 5.15. Distribution of Distribution 5.15. Figure (2007d). Dijkstra From: species. sympatric 148 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA nearest relative inSouthAmericaandwasbelievedtobe are present (see2.2.3;Figure 5.7): genera withdistinctMadagascanandNeotropical affi (Figures 5.14-5.15).Moreover, severalrelict damselfl Chlorocnemis groups suchasCalopterygidae,Chlorocyphidae and has Africa’s richestodonatefauna:typicalrainforest This region from south-eastNigeriatonorth-westAngola 5.3.2.3 LowerGuinea central rainforest blocks. NigeriathatseparatesAfrica’sand western and western endemic togalleryswampforest inthesavannahofBenin respectively. material fromTogo western Côted’Ivoire, andnorthern and potentiallyconspicuousspeciesknownonlyfrom type jejuna endemics, andasurveyisurgently required. Guinea mayharbourvitalpopulationsoftheseandother Nimba. Theunexplored FoutaDjallonMountainsinwest typical oftheUpperGuineanhighlands,centred onMt. infumosa Sierra LeoneandLiberia. closely related (Dijkstra unpubl.). The genus remains endemictothe Lower Guinea. (Dijkstraunpubl.). Thegenusremains closely related and morphological comparisonssuggestthevulnerable (a) Until recently, thethreatened Two deforestation and mining pose an increasing threat to its survival. toitssurvival. threat andminingposeanincreasing deforestation (b) a (b) (a) (b) (a) S. fumosa Sapho (CR)and ) and species endemic to western Africa.While(a) speciesendemictowestern (DD) is restricted to rocky streams. As a consequence it is largely limited to the Upper Guinean highlands, where Asaconsequenceitislargelylimitedtothe UpperGuineanhighlands,where torockystreams. (DD) isrestricted Ceriagrion citrinum are uptotwiceasrichhere aselsewhere Phyllomacromia funicularioides Zygonychidium gracile Africocypha lacuselephantum Sapho fumosa (VU)ispeculiarinbeing Pentaphlebia (CR)are distinctive (NT; = Chlorocypha (NT)appear hasits S. ciliata Umma “Chlorocypha” centripunctata nities VU from Cameroon was considered the only species of its genus. Genetic theonlyspeciesof itsgenus.Genetic VUfromCameroon wasconsidered y © P. NASKRECKI(a)ANDW. KAPPES(b) (LC) can be found on almost any running water with a bit of forest, (LC) canbefoundonalmostanyrunningwaterwithabitofforest, only non-Americanperilestid(see5.2.2.3;Figure 5.7).Other 2011). Thedefi deforestation, especiallyinNigeria(Dijkstra highlands. Severalare globallythreatened dueto species occursinandaroundCameroon thesouthern The greatest concentrationofrelicts andrestricted range Africa(Figuresand southern 5.18-5.19). the mostdistinctlyAfro-montane characteroutsideeastern speciosus Diastatomma tricolor rufi Guinea endemicsinclude species (Dijkstraunpubl.).OthercharacteristicLower Nesolestes despite theproximity ofbothtotheMadagascangenus Nesolestes nigeriensis west RepublicofCongo.Theformeriscloselyrelated to occur from thesehighlandstotheMayombeHillsofsouth- Neurolestes trinervis west RepublicofCongo(Mézière andLambret unpubl.). third taxonwasdiscovered insouth-eastGabonandnorth- endemic totheCameroon highlandswithtwospeciesuntila tibia , Chlorocnemis contraria . Furthermore, theLowerGuineahighlandshave , probably represents asecond © K.SCHÜTTE (a)ANDC.VANAPPELGHEM (b) ning speciesis and (b) an unnamed species from Gabon are and(b)anunnamed speciesfromGabonare , and Phyllomacromia insignis from theCameroon highlands,which Chlorocypha gracilis Stenocnemis pachystigma Nubiolestes diotima , Platycnemis rufi and , et al. Neurolestes Platycypha (VU), the Zygonyx 2010; both pes ,

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 149 Lokia (c) LC, of © K.-D.B. DIJKSTRA Neurogomphus martininus Neurogomphus (LC) and (c) (a) (b) Blackwater swamps and rivers (a) are characteristic of the Congo Basin and are home to endemic species like (b) erythromelas which a larval skin is shown.

, , , , T. P. in was (EN), (DD) 2011). Porpax (CR), (CR), and , Nesolestes Nesolestes G. quarrei Elattoneura Elattoneura are known are Elattoneura Trithemis et al. nds placing Chlorocypha Chlorocypha Nesciothemis Pentaphlebia , T. hartwigi T. , (DD), Trithetrum navasi Trithetrum and (VU), (VU), Sapho puella (VU), ned to the Congo (CR). Until recently, (CR). Until recently, Zygonyx . Remarkably, many . Remarkably, Notogomphus mooreiNotogomphus , and Phyllomacromia caneri Phyllomacromia Trithemis hartwigi Trithemis (DD), da Trithemis aconita Trithemis 2011): the Malgassophlebia and Mesocnemis (LC), , , , (LC), I. regisalberti Zygonyx Pentaphlebia gamblesi Pentaphlebia T. bifi T. et al. U. purpurea ned to standing waters, possibly ned to and and (NT) and tzgeraldi and C. neptunus C. neptunus Gomphidia gamblesi (EN), Mesocnemis Africocypha . The research also produced new species also produced . The research N. fi Chlorocnemis eisentrauti Chlorocnemis , , which had been photographed 26 years Pseudagrion risi Pseudagrion (VU), Lestinogomphus , and T. basitincta T. Urothemis Africocypha lacuselephantum Africocypha (CR), , Neodythemis takamandensis , (LC), cally crater lakes, in Bioko and adjacent Cameroon, Bioko and adjacent Cameroon, cally crater lakes, in Tragogomphus aurivilii Tragogomphus (VU), Urothemis T. congoense T. Pentaphlebia pruinosa (LC), congolica Pseudagrion (LC) and Azuragrion buchholzi thought to be confi were specifi Gabon, while but both occur in south-east nigeriensis also found in south-western Cameroon and photographed also found in south-western Cameroon pers. comm.). Similar fi in east D.R.C. (Windig The area, likely. danger are species outside immediate harbour undiscovered may also than any other, more species. threatened of the southern Guinea, from The rainforest Lower Gabon to south-west southern through Cameroon the odonatologically least known, Republic of Congo, are equally rich (Dijkstra although it is probably Ictinogomphus fraseri This is demonstrated by recent research by N. Mézière in by N. Mézière research This is demonstrated by recent south-east Gabon: in two years, Haut Ogooué Province, of about 200 x 100km at 400-700m altitude, in an area Among these are over 180 species have been recorded. as well as range extensions of rediscoveries, numerous confi species thought to be (largely) further north Basin or to savannah and woodland much complexity is or south. Illustrative of local diversity and forest-savannah that many north-south, west-east and unknown to overlap, meet species pairs, which were for example, here, characteristic endemics are are endemics characteristic stahli centripunctata nigeriensis Umma mesumbei Umma mesumbei of and north of Yaoundé (750km north), and the north of Yaoundé is the (625km north-west). Most extreme south Cameroon new earlier 1,500km east in D.R.C. (Windig pers. comm.). This rich demonstrates that our knowledge of this incredibly studied. are fauna will rapidly expand once further areas 5.3.2.4 Congo Basin woodlands and rivers, swamps, forests, Ever-shifting an enormous mosaic savannahs in the Congo Basin create heart of prime odonate habitat in time and space. Africa’s between the forests has always been on a crossroads, to the west and east in wet periods, and between the Trithemis (Dijkstra not localised are for example, was found about 350-400km south-west in the north-western pers. Republic of Congo (Lambret comm.), while the 150 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA dark grey line:route Congo2010expedition.Adaptedfrom: Dijkstra River watershed;dashedellipse:approximate extent of“cuvettecentrale”,defi circles: siteswithreasonable odonatologicaldata,noneofwhichare CBFPpriorities;palegrey area: Congo priority according toCongoBasinForest Partnership,noneofwhichhavebeensurveyedforOdonata;fi Figure 5.16.OdonatologicalexplorationoftheCongo Basin.Legend–opencircles: sitesofhighconservation aphrodite of thebasin,and largely confi Africa (Dijkstra data largely comefrom the RoyalMuseumofCentral sampled mostlyinthe1930sto 1960s(Figure 5.16).These our knowledgeislimitedtoahandful ofperipheralsites, true forOdonata(Dijkstra2006c; 2007d).Unfortunately high endemismanddiversity, whichcouldbeespecially diversifi that thebasinisan“evolutionarywhirlpool”ofspecies the basinandalongitsrivers.Kingdon(1989)postulated forest speciescouldsurviveinrefuges west andeastof savannahs tothenorthandsouthindriertimes,when Prodasineura odzalae cation, conservationanddispersal,leadingto , Elattoneura centrafricana et al. 2011).Speciesfoundinalarge part , Pseudagrion simplicilaminatum ned toit,are , E. vrijdaghi Chlorocypha , .P. , martininus Zygonyx regisalberti and in theCongoBasin bylarger anddarkerforms: widespread non-forest groups (see5.3.1)are represented like huge rivers,oratleastwiththe associatedswampforest, Zygonoides occidentis (= maesi regisalberti thenartum apicalis erythromelas Congothemis Agriocnemis stygia Z. fuelleborni , (= P. schoutedeni , et al. , , Porpacithemis trithemoides N. uelensis Diastatomma multilineatum , Phyllogomphus annulus Porpax garambensis (2011). ) replace longistyla . Speciessuchas ned aspartofbasinbelow500m; are (near)endemicto theregion’s and , , Paragomphus acuminatus P. seydeli Hadrothemis vrijdaghi , Z. occidentis Trithetrum congoense , P. sentipes , , ), , Lokia circe onopen rivers T. congolica Phyllomacromia Neurogomphus Ictinogomphus , lled Trithemis Z. fraseri . Several , and and , , L. T. T.

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 151

, rst with et al. et al. species, Aciagrion Ceriagrion Platycypha ; , (both DD), and T. congoense T. , for example, had Gynacantha Prodasineura . Discovery of Z. fuelleborni (2006b). (2006b). Pseudagrion malagasoides et al. Platycypha eliseva and were found, the second and ﬁ found, the were and ; triangles: ; triangles: Paragomphus and Chlorocypha pyriformosa Ceriagrion Z. lachesis , the only species of the recently erected genus erected , the only species of the recently and Elattoneura morini , this seemingly monotonous forested plain crossed by plain crossed this seemingly monotonous forested 86 species within 13 days in a countless rivers, recording the widespread; Lokutu. Only 28% were around small area than 125 species. fauna was estimated to number more two conspicuous new species, The survey produced Mesocnemis saralisa range extensions over thousands of kilometres. ignitum not been found outside western the former Africa before, Ghana (Dijkstra not even since its description from 2011). An expedition in April to June 2010 on the Congo, 5.16), yielded Kisangani (Figure from going downstream many additions: including prior data, 171 species have new Two of the river. been found along this 350km stretch species of male specimens of two unnamed species of new and probable Notogomphus picta not Batéké plateau and western shows that these are assumed (Dijkstra African endemics, as recently 2010; 2011). The extensive sympatry of navasi T.

, Chlorocypha ; shading: inferred ranges. From: Dijkstra Dijkstra From: ranges. inferred ; shading: species. Legend – squares: – squares: Legend species. (Dijkstra 2007a). Rhyothemis splendens Gynacantha manderica , Neurogomphus vicinus ed in the Congo Basin. ed in the Congo Basin. , Z. occidentis Zygonoides T. congolica T. P. maynei P. , , seem associated with large rivers. , seem associated with large . The most notable hiatuses are . The most notable hiatuses are ; circles: ; circles: occupy half-shaded habitats in most Elattoneura morini Z. fraseri , Z. occidentis Trithemis fumosa Trithemis Trithemis aconita Trithemis ciently from the periphery to diverge genetically. the periphery to diverge ciently from rst recent impression of the remarkable diversity of of the remarkable impression rst recent and large streams in the dry northern and eastern streams belts and large 5.17). of continental Africa (Figure and of tropical Africa, excluding only the driest deserts and Africa, of tropical by a in the basin former is represented The densest forests. form that merits species status (Dijkstra 2005c; see below), by the latter is replaced Both, like of diversity discussed, no threatened Unlike other centres or NT species have been identifi pristine, but also lack data, parts of the basin are Large DD – for example, and quite a few species are ghesquierei Forest expansions and contractions may have enclosed expansions Forest matrix, isolating them these populations within the forest suffi Paragomphus interruptus and on the eastern slope (see 5.3.4.1) and in the “Cuvette of the central basin area Centrale”, the almost circular 5.16). Dijkstra (2007b, 2008) obtained below 500m (Figure a fi diamonds: diamonds: Figure 5.17. Distribution of Distribution 5.17. Figure 152 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA subgroup (grey –R: N: unnamedspecies;X: (e: Figure 5.18.DistributionofAfrotropical (a) A. e.ellioti ©

E.R. NIELSEN(a)ANDK.-D.B.DIJKSTRA(b) Notogomphus moorei characteristic Afro-montane genera. characteristic Afro-montanegenera. ; u: (a) A. e.usambarica Aeshna minuscula A. rileyi A. moori ; S: A. subpupillata (LC) represent (LC) represent ; Y: (LC) and (b) (LC)and(b) ; m: A. yemenensis A. minuscula Aeshna ; “?”:Angolarecord, speciesuncertain).From: Dijkstra(2007d). species.Legend–ﬁ ; “?”:Sudanrecord, speciesuncertain)andsouthern ); outlines: rileyi -group (black–M: withnorthern lled circles: A. scotias ; shading: A. meruensis ellioti -group (b) ; CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 153

also Proischnura Notogomphus Notogomphus Pseudagrion and (Figure 5.18), (Figure (Figure 5.21) and to some degree to some degree 5.21) and (Figure Aeshna , and the A-group of and the A-group Pseudagrion spernatum Atoconeura (Figure 5.20) most clearly demonstrate an Afro- 5.20) most (Figure , few taxa inhabit the total expanse of this Proischnura Africallagma uence (5.3.5.1) on the periphery (Figure 5.10). on the periphery (Figure uence (5.3.5.1) (Figure 5.19), (Figure Platycypha montane distribution. For instance, of 19 species, 16 occur east of the Congo Basin and four to the west. diversity and endemism. All show marked Afro-montane abundance above about their greatest have these groups 1,000m altitude. The transition to montane habitat on the equator is at 1,500-1,800m (Dowsett 1986) and the upper 2,800-3,300m (de Jong and Congdon at limit of forest 1993), but a truly montane fauna, with species occurring exclusively above 2,000 or even 3,000m is virtually absent complexity is the (see 5.3.3.4). Adding to the Afro-montane (see 5.3.4.1-2) and even insular Guineo-Congolian strong infl here than in western diversity here and central Africa – national – but regional generally lies between 100 and 200 species Except for some ubiquitous highland endemism is greater. species like subfurcata The geographically and climatologically diverse region. genera . Figures indicate the number of allopatric species; local peaks in local peaks species; allopatric number of the indicate . Figures ed an “eastern Notogomphus (Dijkstra and Pilgrim 2007) is also notable. (Dijkstra and Pilgrim 2007) is also notable. Trithetrum the southernDespite these efforts, and western Cuvette, of north-east Republic of including the swamp forests unexplored. entirely Congo, remain fauna 5.3.3 Afro-montane 5.3.3.1 Overview Eastern and southern but Africa harbour diverse of higher to areas restricted fragmented forests, elevations. White (1981) at greater especially precipitation, archipelago” characterised these as an “Afro-montane Dowsett- because of their isolated and dispersed nature. (2001) further identifi and Dowsett Lemaire The term Afro-montane forests. of coastal archipelago” fauna: “montane” forest is perhaps not apt for the entire descends to sea-level in South Africa, while localised and swamps at moderate inhabitants of coastal forests included. Nonetheless, almost all of also altitudes are where Africa above 1,000m is included within the region occurs, and most of this region the fauna discussed here lower is also above that altitude. Species numbers are Figure 5.19. Distribution of Distribution 5.19. Figure from: Adapted sites. as separate given are records Guineo-Congolian Scattered shaded darker. are diversity in prep.). Dijkstra (2007d; 154 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA logical base for the eastern radiation. Fourspecies have logical base fortheeastern With apresumed ancestor, western the AlbertineRiftisa locality alsohas theonlystandof spray zones(White1981).Indeed, Ghana’s sole plants similarlysurviveinspecialised habitats,suchas the baseofhighlands.Lowland populationsofmontane has amontanecharacter, occurringnearwaterfallsat picta Figure 5.20.Distributionof intervening habitatislow-lying. Nonetheless, lowland representative isalsoeasier, simplybecausemost at muchhigheraltitudesinthewesttoo.Dispersalof westwards ratherthaneastwards, itmight beexpected eastern “basal” positionsuggestsitmaybeancestraltothe A. luxata inhabits streams above1,000m,allrecords ofthewestern 2006b; 2007d;Figure 5.21).Whilethegenusmainly illustrate thehistoryofAfrica’s highlanddragonfl isolation ofthehighlands.Thegenus vicissitudes atgreater altitude,aswelltheyouthand is related tothemore severe impact ofclimatological The relative impoverishmentoftheAfro-montane fauna population of P. inyangae ; r: Atoconeura liebelowit.Itsphylogeneticallyandecologically P. rufi ; p: tibia P. caligata P. pinheyi ; e: radiation.Hadthegenusdispersed P. eliseva . From: Dijkstra(2008). ); squares: Platycypha ); fi Cyathea lled circles: sub-montanerelict species(a: P. lacustris species.Legend–opencircles: lowlandrelict species(a: Atoconeura tree-fernsthere. ies (Dijkstra A. luxata A. luxata ; shading: may

P. caligata relict species (Samways1999, 2008).Almost one-fi centres ofendemism,harbouringthe greatest numberof The continent’s end isoneofitsmostsignifi southern 5.3.3.2 SouthAfrica periods andunreachable whenhabitatswere suitable. South Africanhighveldwasmaybe uninhabitableincooler and suitablehabitat:itmaystill be foundinAngola,butthe notably absentintwohighlandregions withhighendemism habitat, withanaltitudinalupward tendency. followed byrapidadaptationanddispersalwithinthenew Atoconeura with thephylogeneticpositionof and mostofitsspeciesspread outfrom Kenya, conforming Kenya ameliorated.Ethiopianmontaneforests are recent, that expandedfromArc thebenignEastern asconditionsin probably developedfrom anancestornear by dispersalacross montanesteppingstones. most proximal geographically, suggestingexpansion recalling a“ringspecies”:theclosestrelatives are also interlocking rangesaround thedrynorthofTanzania, ; triangle:morphologicallydeviantlacustrine historyrefl P. amboniensis ects phasesof habitatopeningup, A. aethiopica ; f: .fi P. P. auripes tzsimonsi Atoconeura A. biordinata . Altogether, A. kenya ; p: fth of fth ; i: cant .P. is

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 155

, , et al. Allocnemis Platycypha , Ceratogomphus (LC) represent the , represents the last represents (Figures 5.14, 5.18, 5.14, 5.18, (Figures A. kenya Proischnura (b) species diverged around 60 around species diverged Metacnemis

, , Syncordulia Orthetrum (LC) and and Syncordulia (Figures 5.7-5.8), and genera (or species 5.7-5.8), (Figures Pseudagrion Ecchlorolestes Aeshna , , inhabits cold and nutrient-poor streams almost inhabits cold and nutrient-poor streams , © N. MÉZIÈRE (a) AND A. CORDERO RIVERA (b) ora. Cold upwelling off the Cape coast began to upwelling off ora. Cold Syncordulia gure approaching one-third if peripheral tropical if peripheral tropical one-third approaching gure Atoconeura luxata (b) (a) 5.20). Occurring in the most remote corner 5.20). Occurring in the most remote of a continent history of isolation, it is tempting to link with a prolonged odonates with that of relict the history of South Africa’s (“fynbos”) they inhabit, which shrubland the sclerophyll is characteristic of the Cape Floristic Region. Ware (2009) estimated million years ago. simultaneously with important radiations of Cape fl block summer rainfall 10-8 million years ago, fynbos only 5-3 million years ago. forest warm temperate replacing Syncordulia devoid of more widespread Afrotropical Odonata. This Afrotropical widespread devoid of more that suggests environment vestiges of an older (>60 million years ago) Afro-montane better assemblage, surviving in a habitat to which they are competition with, modern adapted than, and safe from Elattoneura (a) ecological and morphological extremes of an Afro-montane radiation. endemic, are 160 species approximately South Africa’s a fi split excluded. The 30 or so endemics are species are (near) endemic genera about equally between the “relict” Chlorolestes and character: Afro-montane with an groups) Africallagma A. (Ethiopia), (Ethiopia), A. pseudeudoxia (Kivu to western Kenya), A. aethiopica A. eudoxia (southern and to Zimbabwe Katanga . The phylogeny of the species is of the species phylogeny . The (northern Tanzania to southern to Sudan), (northern Tanzania (Sierra Leone to Gabon) and (Sierra Leone to Gabon) southern Kenya), A. kenya A. biordinata luxata (northern Dijkstra Zambia to western Uganda). From: (2006b; 2007d). Figure 5.21. Biogeographic hypothesis for for hypothesis Biogeographic 5.21. Figure Atoconeura altitudinal ranges. on their observed superimposed of species is Distribution 156 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA (DD) are the onlyonesavailable. (DD) are Indeed ourregional knowledgeisverypoorandonly distinct uplandfaunamaybeduetoinsuffi affi The fewAngolanendemicsknownhaveGuineo-Congolian 5.3.3.3 Angola 5.4.2.4). trees andfi in Africatwoofthemajorregional threats, invasivealien 1977 andmaybeextinctorhighlythreatened. Moreover, (VU). S. serendipator Syncordulia gracilis (EN), Proischnura polychromatica E. peringueyi Chlorolestes apricans and oftenshortfl (Samways 2006a),manyofthemwithverysmallranges in Africa,withalmosthalfoftheendemicsthreatened second-largest concentrationofgloballyimperilledspecies regard totheirstatusandthreat. Theregion holdsthe that theSouthAfricanendemicsalsostandapartwith habitat isalastrefuge from extinction,itisunsurprising tropical species(Dijkstra Madagascar andPemba(Dijkstraunpubl.). (a) Angola’s of (b) poorlyknownandthesepictures very endemics are Both speciesof (a) (a) nities (see5.3.1.1),buttheapparent absenceofa M. valida Orthetrum rubens P. newtoni sh, are virtuallyuniquetoSouthAfrica(see (EN) is related tothetropicalmainlandgenus (EN) isrelated (VU), Metacnemis (NEasnewlydescribed),and ight periods(SamwaysandGrant2006a): (VU), Metacnemis angusta (VU), (EN), (DD)hasnotbeenrecorded since Ceratogomphus triceraticus S. legator et al. inhabit tiny and encroached ranges in South Africa. While unpublished genetic research showed inhabittinyandencroachedrangesinSouthAfrica.Whileunpublishedgeneticresearch (CR), Ecchlorolestes nylephtha

(b) bivittatus Pseudagrion angolense (all DD): not beenrecorded since,mainlyoccurinthehighlands other speciesdescribedfrom Angola,manyofwhichhave from thenominate(Dijkstraunpubl.).Itisalsolikelythat to (see 5.2.2.3;Figure 5.9). (VU) isoneofAfrica’s greatest biogeographicanomalies smallest rangesinAfricanOdonata,while completely isolatedtaxonomicallyandhaveamongthe of which than fourmonotypicgeneraare endemicandthreatened, north-east SouthAfrica,andtheadjacentcoast.Nofewer species occursinthemountainsstretching from Kenyato Africa’s foremost concentrationofthreatened orNT MountainsandCoast 5.3.3.4 Eastern and species eastern and thelocalsubspecies For example,anunnamed research willyieldnew, possiblythreatened, endemics. Umma femina two speciesare currently assessedasthreatened orNT: Chlorocypha crocea (b) Tragogomphus Micromacromiaﬂ (b) M. angusta , Amanipodagrion Onychogomphus rossii Chlorocypha rubriventris (VU)and Platycypha caligata andmayrepresent aGuineo-Congolian (VU)iscloserto (NT)and(a) ava , Chlorocypha crocea . Nepogomphoides P. dundoense and angolensis Elattoneura Oreocnemis Oreocnemis Pseudagrion angolense , Paragomphus machadoi Platycnemis is geneticallydistinct , of thewidespread isalready known , Aciagrion rarum Lestinogomphus (VU)issimilar (both CR)are Coryphagrion (NT).Further of

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 157

(VU) (VU) Oreocnemis Chlorolestes ) extends to RG s (CR) is found (VU Teinobasis alluaudi Teinobasis (VU) and (CR) only inhabits Chlorocnemis montana Africallagma cuneistigma (NE, but considered as a (NE, but considered Coryphagrion grandis Pseudagrion vumbaense C. maccleeryi Platycnemis pembipe Teinobasis alluaudi Teinobasis Chlorolestes elegans Chlorolestes Platycypha inyangae (VU), (b) (VU), the only African synlestid outside South (VU), the only African synlestid outside (CR) is endemic to Mulanje, the highest mountain (CR) is endemic to Mulanje, the highest mountain (EN) occurs in mountains on both sides of the Tanzania- (EN) occurs in mountains on both sides while Malawi border, 5.14). Mount Ntchisi in central Malawi (Figure phoenix with a 2,000m and the Drakensberg, between Kilimanjaro the from high and 24km wide plateau rising almost vertically 700m high plains of southern Malawi. The species’ 50km² vulnerable to bauxite mining. range is entirely elegans but population here, Africa (see 5.3.3.2), also has a strong in the north-east of South extends to the Soutpansberg Africa and Mount Namuli in northern Mozambique. In the local mountains of eastern it occurs with three Zimbabwe endemics: species by Dijkstra (2008)), and (VU) in the Chimanimani and Mountains. in the Vumba On the coast, the endemic while occurs in Kenya and Tanzania, southern (EN) extends from Kenya inland to northern is LC if its Madagascan however, Malawi. The latter, and highly threatened (both CR). and highly threatened (b)

Umma declivium is (a)

(LC) (VU) (VU) to (b) (c) P. bicoerulans P. (NT) (Figure 5.14) (NT) (Figure (CR) at 1,500-2,000m, Atoconeura kenya Amanipodagrion gilliesi (VU) occurs both in the are unique to the latter unique to the are Umma declivium © K.-D.B. DIJKSTRA (a), W. TARBOTON (b), A. CORDERO RIVERA (c) (b), A. CORDERO RIVERA TARBOTON © K.-D.B. DIJKSTRA (a), W. (VU) and (EN) at 2,200-2,600m, and only ies from the easternies from highlands: (c) VU. Nepogomphoides stuhlmanni © J. RENOULT (A) AND V. CLAUSNITZER (B) (A) AND V. © J. RENOULT Platycypha auripes Chlorocnemis abbotti Platycypha amboniensis Micromacromia miraculosa (a) (a) up to and occasionally over 3,000m. being above the most “alpine” African odonate, typically and 2,500m and not found below 2,000m (Clausnitzer Dijkstra 2005b). mountains Of species occurring mainly in the Eastern Arc of Tanzania, extends to southern Kenya, northern Malawi, and to northern Mozambique and southern Of the Malawi. true endemics, Udzungwas and Usambaras, but and the adjacent mainland, Madagascar (where this picture was taken) and the Seychelles, (b) nowhere else on Earth. Pseudagrion bicoerulans Both these threatened damselflies occur in the tiny Ngezi Forest on Pemba, but while (a) Notogomphus maathaiae Pseudagrion bicoerulans relict (see 5.3.2.1). Most endemic species, however, (see 5.3.2.1). Most endemic species, however, relict radiations (see 5.3.3.1). belong to typically Afro-montane highest-living This is, for instance, the case for the in Kenya and north odonate fauna, centred Afrotropical Mounts Elgon, Kenya, Meru, Kilimanjaro around Tanzania Their habitat is being encroached and the Aberdares. to altitudinal is related uphill, hence the level of threat occurrence: Three threatened damselfl 158 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA Paragomphus Pseudagrion are fourendemics.Almost allbelongtogenera( the 14mostwidespread high-altitudespeciesinEthiopia Kenya, Tanzania andUgandaeach(see5.3.3.4).Among In contrast,endemismishigh at12%,versus1-3%for presence ofsuitable habitat(Figures 5.12,5.14-5.15). Micromacromia dominate furthersouth, mostappearing closelyrelated to like Guineo-Congolian forest connection.Forest genera Africa,areUganda towestern theonlysignsofaformer Gynacantha forest species(ClausnitzerandDijkstra2005a).Two The impoverishmentrests mainlyonthepaucityof of 12endemicodonatesconsidered threatened. surviving inastrongly impactedlandscape,withnine comparatively species-poorbutendemic-richfauna, assemblages (see5.3.3.2).LikeSouthAfrica,ithasa trioofmostsensitive Ethiopia completestheeastern 5.3.3.5 Ethiopia 5.22). by Dijkstra is notarecent arrivalthere from Madagascar, asthought a singlestream. Unpublishedgeneticdatasuggeststhatit Tanzanian coast,onlyin2001,andisstillknownfrom pembipes and Seychellespopulationsare considered. (unpubl.). Madagascar seemtoformtheperipheralremains ofanolderAfrotropical radiation.From: DijkstraandStokvis Congolian taxaare mostcloselyrelated totheAsians,whilespeciesinCape,Pemba,Comoros and legged genetic data,issuperimposedonthegeographicranges.Thespeciesare more closelyrelated totheyellow- Figure 5.22.BiogeographichypothesisforAfrotropical taxaplacedin Chlorocypha Copera (CR)wasdiscovered onPembaisland,off the et al. species,thatotherwiserangefrom central , , Elattoneura Atoconeura and species of southern Asia,thanthewhite-legged speciesofsouthern (2007a),butratheranancientrelict (Figure , Umma Notiothemis , , , Notogomphus Orthetrum Chlorocnemis are absent,despitethe and , , Crenigomphus Trithemis Hadrothemis Platycnemis Aeshna ) that , , , and Ischnura abyssinica immediate conservationmeasures: being sensitiveduetotheirsmall ranges,butnotrequiring half oftheendemicsinhabitwholly altered landscapes, almost completelytoagricultural land.Indeed,abouta change, whichhastransformed thenaturallandscape Odonata mightberelatively toleranttoanthropogenic fromclimatic vicissitudes.Borne suchadversity, Ethiopia’s adapted tomontaneconditionssurvivedthesubsequent expanded intotheEthiopianhighlands.Especiallythose or wetterperiodstropical Africanspeciesmusthave colonisation ofthehighlands.Nonetheless,inwarmer barriers, evenforopen-landspecies,hampering and theescarpmentsofEthiopiaitselfare formidable Sea, desertsofKenyaandSomalia,WhiteNilefl between thedescendingcoldandaridpiedmont.TheRed and wetconditions(likeOdonata)were literallycrushed During coolerperiods,speciesadaptedtorelatively warm to thehighlands’large massoflandatgreat altitude. Climatic fl endemic-rich character(ClausnitzerandDijkstra2005a). history andisolation,explainsitsspecies-poorbut High altitude,strong seasonality, aswellEthiopia’s (Dumont Only species inthehighlandsthere (Figures 5.18-5.19,5.21). LC). Of more are concern those endemics thatinhabit Orthetrum kristenseni Ischnura abyssinica Platycnemis et al. uctuations were probably relatively severe due Platycnemis 2009). speciesofEurasia.TheGuineo- and possiblyhasPalaearctic affi . Thephylogeny, basedon and Pseudagrion kaffi Trithemis ellenbeckii Elattoneura pasquinii num oodplains (allVU), (both nities , CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 159

bwambae (NT) was (NT), and Tetrathemis Tetrathemis C. schmidti Agriocnemis ) , cult to survey, and cult to survey, Chlorocypha hasta (NT), Idomacromia jillianae Neurogomphus wittei N. nyungwe sh and molluscs have . However, in the Albertine . However, (DD), Platycypha pinheyi (DD), Lokia coryndoni Onychogomphus , 2011), but many endemics or C. molindica y species from Lakes Tanganyika Lakes Tanganyika y species from (= avifrons et al. was discovered on reedy beaches on reedy was discovered T. denticauda T. (NT), (DD). Most of these are only or mainly (DD). Most of these are and (NT) is known only from inside papyrus swamps (NT) is known only from A. pseudeudoxia Pseudagrion , Tragogomphus C. jacksoni Notogomphus fl Neodythemis munyaga (DD), (DD), ruwensoriensis T. because the Uganda, probably south-west known from western (Congolese) slope of the rift is almost completely unstudied (Dijkstra too. must occur there new species related palaeforma but diffi it is numerous in Uganda, where widespread. it too may be more such as fi While aquatic groups Lakes, only Great “explosive” endemism in the region’s a handful of dragonfl sea” endemics: and Malawi may qualify as “freshwater a new of the former near Kigoma, and shores, on rocky numerous (LC) may also be endemic (Dijkstra 2009). This paucity (VU), eudoxia corduliformis or found that are localised species are Highlands, more or (mostly) NT: may be threatened (VU), (EN) are endemics threatened Ethiopian by habitat destruction.

N. C. , and (EN), Atoconeura , Paragomphus Paragomphus Notogomphus , , (VU). The habitats Neodythemis (VU), , Chlorocnemis pauli Notogomphus cottarellii Notogomphus cottarellii , Crenigomphus abyssinicus . Many of the species can C. trifaria Notogomphus cottarellii Chlorocnemis , (VU) and (b) , Phyllomacromia sylvatica (both VU), are unknown. (both VU), are Atoconeura , Onychogomphus styx (DD), Atoconeura aethiopica , Pseudagrion guichardi and , as well as in the more montane genera , as well as in the more Africallagma pseudelongatum Chlorocypha P. viridior P. , , N. lujai (EN), and , C. denticulatus Chlorocypha tenuis Elattoneura pasquinii (b) (a) of the two remaining endemics, of the two remaining ruppeli and 5.3.4 Complex faunas 5.3.4.1 Albertine Rift of central Africa slopes up to meet the The low-lying forest eastern across Africa that stretches huge elevated area along the lie here, highest forests 5.11). Africa’s (Figure western (Albertine) rift in east D.R.C. and adjacent Uganda, widespread Aside from Rwanda, Burundi and Tanzania. (see 5.3.3.1), these highlands have species Afro-montane fauna, with endemism in genera a characteristic forest such as crenigomphoides forest streams: streams: forest superba leroyi lacustris Tetrathemis Notogomphus hardly be called “Albertine”, as they range to central, north- be called “Albertine”, hardly east or southern D.R.C. and/or westernExamples Kenya. are (a) © K.-D.B. DIJKSTRA 160 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA The rocky shores ofLakeTanganyikaThe rockyshores theuniquehabitatof(b) (a)are but hascompletelydifferent coloration.The of slow-fl Pseudagrion lakeshore populationsfrom surrounding habitats.Thenew while itscapacityforfl many lakenichesinaccessibleforlarvaldevelopment, can beexplainedbytheadult,whoselinktolandmakes fauna must bemonitored. and itsuniqueecological andevolutionarycharacter, this seven are assessedasNearThreatened, andowingtothis None ofitsendemicsare asyetgloballythreatened, but recent speciationcentre forOdonata(Dijkstra2007d). on thisgeologicalcomplexity, the region isprobably a Larkin 2001).WithAfrica’s climatic historysuperimposed drained headwatersthatnowfeed theCongo(Moore and of recombination: theZambeziandevenLimpopoonce drainages. Theregion’s rivershaveacomplexhistory is nowsplitacross theCongo,OkavangoandZambezi which drainedintotheMakgadikgadiPaninpast,but delimited bythePaleo-Chambeshisystem(Figure 5.12), The area liesintheheartofZambezianbiomeandis Aciagrion 16 are inthree ofthemostsuccessfulAfrotropical genera: Afrotropical odonates,withabout55endemics, ofwhich Africa. About325speciesoccur, anestimated40% of all one ofthemostimportantcentres ofodonatediversityin through AngolaandZambiatonorthBotswana,as eastern Pinhey (1978)already identifi 5.3.4.2 Zambezia reinforced bysexualselection. coloration, whichinthisvisuallyorientedorder maybe from riverinefoundersisfollowedbyslightdivergence in only occurringwheninitialseparationofalakepopulation endemic Odonatamaybearecent phenomenon,possibly frompattern itsriverinecounterpart.Theevolutionoflake caligata and anotheronLakeMalawiclassifi a (b) (a) (Dijkstra2005b),differs onlymarginally incolour Africa, owing andstagnanthabitatsineastern , Pseudagrion isstructurallyidenticaltoacommonspecies and ight inhibitsgeneticisolationof ed thearea from Katanga, Trithemis ed asaformof (Kippingunpubl.). Platycypha .P. , (NT), restricted African speciesare found inrunning waters, drowned (LakeMweru).Whilethevast majorityofrange- once existedin basins nowdrier(Etosha,Kafue, Lufi “piracy” (seeabove)andalluvial depositssuggestthese systems. Thelattermaybea transientfeature ofriver swamps suchastheBangweulu, OkavangoandUpemba depressions, oftenassociated withgalleryforest) tohuge habitats. Theyrangefrom thenumerous “dambos”(wet Especially notableistheregional abundanceofswampy streams, ditchesandpoolsinopengrassland. only dependsongalleryforests, butalsoonbogs,rivers, (1984), anddemonstratedthattheregion’s richness not endemics, whichwere unrecorded since1972byPinhey A. Güntherin2010confi the headwatersofZambeziRiver, byJ.Kipping and Fieldwork intheIkelengePedicleofnorthZambia,with Lokia ellioti Phyllogomphus schoutedeni kitchingmani soror P. symoensii Pseudagrion coeruleipunctum Chlorocypha frigid or “mushitu”(Pinhey1978).Amongtheendemicsare [near] endemicsare known”,knownas“litu”,“muhulu” fringed withthickgalleryforest”, where “agreat many tending toswampformationinparts,andmore orless to large lowlandrivers.Characteristicare “streams, all varietiesoffl along theregion’s ratherhighrelief. Theregion harbours their mainrange,whileAfro-montane taxacandisperse allow Guineo-Congolianspeciestopenetratewellbeyond montane, groundwater andgalleryforests. Thesegalleries landscapes: from grassland,savannahandwoodlandto 1,500m inaltitude,withacorresponding diversityof The region isapatchworkofplainsandplateausvarying Platycypha pinheyi , Chlorocnemis wittei Neurogomphus pallidus , Neodythemis fi (NT), (DD), owing water, from uplandstreams , (NT). C. seydeli Paragomphus cataractae Ceriagrion sakejii rmedthepresence ofmanythe © K.-D.B.DIJKSTRA tzgeraldi , Prodasineura fl , Phyllomacromia unifasciata (DD), (NT), , , and Allocnemis mitwabae P. fi P. Zygonyx atritibiae Onychogomphus sheri , avifacies Diastatomma , P. greeni (NT), (DD), ra) or ra) , . ,

(a)

(b) (a) Umma electa (LC) and (b) Nesciothemis fitzgeraldi (LC) are just two spectacular species discovered during recent fieldwork in north-west Zambia. © J. KIPPING 162 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA species wasfi afterthe of whichwasphotographedabout halfacentry species demonstrated bythisspectacular unnamed(a) Madagascar’s poorlyknown, as very odonatesare The genus (a) ANDC.COHEN(b) (b) (a) and(b) Pinheyagrion rst describedbasedonafemale. Viridithemis viridula hasonlyonespecies, (DD),thefi rst male rst P. angolicum Platycnemis © K.SCHÜTTE (LC),whichonlyoccursintheswampsofZambezia. (near) endemicsinclude restricted-range odonatefaunaofopenswampsinAfrica: species are widespread. Zambeziaharbourstheonly especially inforests andhighlands,mostswamp-dwelling katamborae species, whiletheremaining endemic fi Malgassophlebia radiations of Anax bangweuluensis and Nesciothemis fi Calophlebia aequalis Isomma to endemicgenera( 95% ofZygoptera)are endemic,ofwhichabout80%belong 175 speciesinMadagascar(60% ofAnisopteraandalmost these andtheirlinkstothemainland.Four-fi faunas, andforcompletenessitisrelevant tobriefl The islandseastofAfricaalsohaveAfrotropical odonate 5.3.5.1 IndianOcean 5.3.5 Insularfauna D.R.C.andmostotherswampareas.in southern Zambia,theUpembasystem headwaters innorth-eastern surveyed fordecades.ThesameappliestotheChambeshi vast swampsaround LakeBangweuluhavenotbeen to thebeststudiedinAfrica(Kipping2000;2003a,b) two, butwhiletheodonatefaunaofOkavangobelongs 1984) isexplainedbyformerconnectionsbetweenthe the OkavangoandBangweuluswamps(Pinhey1976; region’s onlyuniquegenus.Thefaunalsimilaritybetween T. brydeni , (NT), Libellulosoma , (DD), Thermorthemis (NT). T. anomala tzgeraldi Nesolestes and Pinheyagrion angolicum Pseudagrion deningi Protolestes Neodythemis (NT), , , , Aciagrion heterosticta Rhyothemis mariposa T. palustris Nesocordulia , , Ictinogomphus dundoensis Platycnemis Viridithemis , Tatocnemis . Theseare largely forest (NE;incl. fth inhabits mostly , , Archaeophlebia , ) ortodistinct fths oftheover P. rufostigma represents the © , Pseudagrion

Paracnemis T. morrisoni J. KIPPING , , Ceriagrion y discuss Trithemis , , , , , ) CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 163

end. dragonfl ies dragonfl Azuragrion Thalassothemis marchali species dragonfl y dragonfl R. KETELAAR (a) AND A. SKINNER (b)

, (km) dist. to land 2007). Allolestes et al. Pseudagrion , (m) of the Mascarenes of the Mascarenes , the Odonata of the max. altitude of Rodrigues is considered invalid. Adapted from: Dijkstra (2010). invalid. Adapted from: of Rodrigues is considered Platycnemis , ) 2 Thalassothemis area area (km Nesolestes Thermorthemis of the Seychelles (ancient continental and and Argiocnemis solitaria Leptocnemis Bioko 2017 3011 34 2 48 0 Príncipe 139 948 210 7 7 1 Socotra 3625 1503 230 6 18 1 São Tomé 857 2024 255 16 14 0 Comoros*Annobón 2023 / 4 21 2361 613 300 340 19 2 22 7 7 0 Cape VerdeRéunion* 4033 / 10 2512 2829 3069 570 600 4 4 14 18 0 2.5 Mauritius* 1865 824 840 7 25 6.5 Seychelles*Rodrigues* 223 / 4 109 905 393 1040 1470 12 2 18 2 5 0 Ascension 97 859 1500 0 0 0 St Helena 122 823 1900 1 1 1 (more recent volcanoes) belong to Coenagrionidae and recent (more Madagascar Libellulidae (see 5.2.2). The Seychelles and Comoros belong to the Madagascan fauna (Dijkstra 2004). Comoros the where Their vulnerability is illustrated in Mayotte, feeder streams, endemic species occur in the very small and human impacts, such as high subject to drought and 2003b). The Mascarene input (Samways detergent summarised Seychelles faunas, whose conservation was (2003a), by Clausnitzer and Martens (2004) and Samways Both island do not contain any Madagascan genera. have two endemic genera, but while groups and 5.7), families (Figure fragments) have been placed in “relict” Coenagriocnemis Nesocordulia Table 5.3. Comparison of Afrotropical islands, sorted by distance to Africa or Madagascar (with asterisk). Cape islands, Comparison of Afrotropical 5.3. Table one and Mauritius share and Seychelles consist several larger islands each. Réunion the Comoros Verde, endemic; open habitats, having close relatives on the mainland (see mainland on the relatives close habitats, having open the island’s on is greatest and diversity Endemism 3.1). wet eastern 2004). While (Dijkstra and Clausnitzer coast with started already of Madagascan Odonata the study in the abandoned was largely research Rambur (1842), cover is forest the meantime, Madagascar’s 1960s. In by 40% (Harper to have decreased estimated (2004) estimated that two-thirds Dijkstra and Clausnitzer possibly are than 100, all endemic) of species (more evaluation. However, immediate and require threatened odonates assessed so far over 70% of 51 Madagascan a single only known from DD, and almost 40% are were the holotype or type series. By collection event, i.e., species have not been found since extrapolation, about 55 still and more discovery, await more discovered, they were becoming known. became extinct before in the western Indian The smaller islands and archipelagos twenty species, of which a Ocean typically have around 5.3). As shown by endemic (Table are quarter to a third endemic species of 164 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA (2007c). Madagascar andtheislandsjusteastofmap’s border, theMascarenes (M)andSeychelles(S).From: Dijkstra immaculifrons Hemicordulia africana Figure 23.Distributionof“insular”Afrotropical Odonata.Legend–blackbottom-leftquarterofcircle: Africa. This ﬁ Tomé (Dijkstra2010), mostofwhichare commonthroughout with seveneach onPríncipeandAnnobón, 14onSão A totalofonly19speciesisknown from theotherislands, it wasconnectedinthepast(Brooks andJackson2001). 5.3). Biokoshares all its specieswithCameroon, towhich Guinea islandsliecomparatively close tothemainland(Table Of thevolcanicAfrotropical islands,theforested Gulfof 5.3.5.2 AtlanticOcean to NT(KetelaarandRiservatoinlitt.2010). potentially byclimatechange,andwilltherefore bereassessed extraction fortheincreasing (tourist)population,aswell the island’s dryclimate,thespeciesissusceptibletowater mountain streams, enclosedinanarea ofonly 450km².Given genus A. granti Despite itsaridity, Socotraharboursoneendemic:the“giant” Africa, probably arrivingrecently from theeast(Figure 5.23). and share Hemicordulia Teinobasis alluaudi Azuragrion (LC)oftheotherwisesmall-bodiedAfro-Arabian gure islow considering the14reported from ; top-right: groups. Each of these also occurs in eastern groups. Eachofthesealsooccursineastern . Itisconﬁ ; bottom-right: , andboththe Teinobasis alluaudi ned toabout10permanent Gynacantha Platycnemis ; enlargedcircles: presence ofthesespeciesorcloserelatives on

bispina speciesofMadagascanradiation;top-left:

Pinhey (1974)sawa common in1990ontheentire island(Gauthierinlitt.2007). one knownis that theislandshaveyettoyieldmore endemics.Theonly endemic genera(see5.3.5.1).Therefore, it seemspossible have muchricherdragonﬂ the more isolated(butalsolarger) MauritiusandRéunion 2008) andthenotablyhighnumberofendemicbirds. Also the more isolateddesert-likeCapeVerdes (Aistleitner from warmer and wetter times, such as 8,000 years ago. from warmer and wettertimes, suchas8,000 yearsago. the Mediterranean, oftensurvivingonlylocally asrelicts Of over800Afrotropical species, fewerthan30reach 5.3.6 Palaearctic fauna recorded in 1963(Pinhey1964). Helena harboured the “giant” but believeditmightbe Gauthier (inlitt.)alsosaw“theenigmaticaeshnid”in1991, on anisolatedislandmightbeexpectedtodistinctive”. São Tomé andremarks that“forest speciesofthesegenera Trithemis nigra Gynacantha Aeshna y faunas,evenharbouring two (CR)from Príncipe,whichwas Sympetrum dilatatum or . The extremely distant St . Theextremely distantSt Heliaeschna Gynacantha species on specieson (EX), last (EX),last et al.

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 165 y

Calopteryx exul (NT) is the only endemic dragonﬂ B. SAMRAOUI

, (NT) and (CR) (VU), (CR), (VU). Libellula Rhyothemis Platycnemis 2010). The Lestes dryas Calopteryx exul (NT), A. isoceles et al. Orthetrum sabina and , (CR), in north-east Algeria. (EN), Coenagrion mercuriale Oxygastra curtisii Pyrrhosoma nymphula ), number just over 50. Lindenia tetraphylla Cordulegaster princeps Coenagrion caerulescens (NT), Sympetrum sanguineum (NT), A. cyanea (NT), which is EN in Europe. (NT), which is EN in Europe. (EN) and I. fountaineae (DD). , (both LC), are NT and VU in Europe and VU in Europe NT (both LC), are (VU), and Cordulegaster boltonii Calopteryx virgo Ischnura saharensis (VU), , Urothemis edwardsii nis Selysiothemis nigra (VU) and , (NT), and have wider ecological and geographic ranges, as have wider ecological and geographic ranges, Ischnura evansi Coenagrion scitulum Gomphus simillimus Lestes numidicus Aeshna affi Gomphus lucasii (a) (b) Although the African populations of (EN), or NT in (parts threatened these species are marginal, are of) their Eurasian range too (Kalkman the western is true for reverse Mediterranean endemic Onychogomphus costae Two more such endemics, more Two in the Maghreb The populations of these three respectively. thus globally important. are quadrimaculata Paragomphus sinaiticus Orthetrum nitidinerve (NT), Boyeria irene O. ransonnetii that south of the Sahara.Thus the fauna stands apart from connected to theEgypt is a notable exception, having been while the hyper- continuously by the Nile Valley, Afrotropics extirpated mostarid period 2,500-2,000 years ago probably Boudot 2010).Eurasian elements (Dumont 1980; Dijkstra and in Europe: All endemics have close relatives of global conservation concern. and are inhabit clean streams Enallagma deserti (EN), subdilatata might impacted in north Africa, severely Natural habitats are due to high human populations and shortage of water. threatened by species that are The severity is expressed populations in or NT in Africa, while they have strong (parts of) Eurasia: (VU), semihyalina by Palaearctic North Africa is inhabited predominantly Asia from species, which together with a few extending (e.g., Consequently, several are Regionally Extinct, or nearly Regionally are several Consequently, Regionally Extinct in northern Africa, for example, (a) Oued Seybouse is the last known site in Algeria for the endangered(a) Oued Seybouse is endemic (b; foreground) North African (EN). Here a male perches with the West Mediterranean endemic Mediterranean (EN). Here West a male perches with the 166 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA of regional endemics insuffi regional lackofknowledgeishighand variable,with16.7% data-defi endemic species)are threatened Africa.Overall innorthern respectively, whileasmany as33.3%(twoofonlysixregionally 20.4% and24.1%ofregional endemicsare threatened, Africa,forexample, andsouthern (see 5.4.1.2):inwestern Threatened (seetable 5.4),withstrong regional differences 9.2% are considered globallythreatened and3.1%Near 88% are endemictotheAfricancontinent.Ofendemics, Slightly more than700specieswere assessed, ofwhich 5.4.1.1 RedListassessmentofAfricandragonfl 5.4.1 Status 5.4 Conservation species across mainlandcontinentalAfrica.Species richness=numberofspeciesperriver/lakesub-catchment. Figure 5.24.Thedistributionof(a)globallythreatened andNearThreatened, and(b)DataDefi (a) ciency fortheAfricanendemics liesat14.9%,but ciently known in thenorth, ies Category represents theirglobal RedListstatus. For allspeciesthat are endemictoAfrica their regional RedList IUCN RedListCategoryataregional scaleinAfrica. Table 5.4.ThenumberofOdonataspeciesineach oa pce 0 621 452 93 705 19 516 95 25 15 Total species 32 Least Concern 18 10 Data Defi cient Near Threatened Vulnerable 37 Endangered 14 Critically Endangered IUCN RedListCategory of species Number cient Odonata Number of endemic species CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 167 ve more NT) and in Angola (one Vulnerable and one NT) NT) and in Angola (one Vulnerable ve more highlands along the Albertine Rift (only two threatened but highlands along the Albertine Rift (only two threatened fi seem less sensitive because habitat destruction is less, The is also considerably poorer. knowledge but regional Katanga to northern Botswana is known for from region its swamp endemism. While none of its endemics are west, most In the assessed as NT. are seven threatened, also inhabitants of highland forests, are species threatened of endemism. in the Lower and Upper Guinea centres concentrated in and and NT species are Eleven threatened border, on the Cameroon-Nigeria the mountains around scattered are one is endemic to Príncipe, and 11 more westernfrom Nigeria to Guinea, though concentrated in the far west. Due to high human populations and shortage impacted in severely natural habitats are of water, only because endemism is low, northern Africa. However, of the The severity globally threatened. species are three by the 17 species is expressed local situation, however, (b) 29.6% in the south, 35.2% in the west, and 41.8% in central Africa. Regional endemism is rather uniform in Afrotropical but with only Africa, lying at just above 20% throughout, 7.4% endemic to the northern with its many Africa region The main cause of these differences, species. Palaearctic level of knowledge in central Africa, is the lower aside from the lower human impact there. species 5.4.1.2 Distribution of threatened or Near globally threatened The 79 African species that are very localised, the vast majority being (NT) are Threatened 5.24a). Seventeen (Figure associated with highland forest in the mountains and coast stretching found species are north-east of South Africa. Fourteen Kenya to the from almost half the occur in South Africa, representing endemics. Ethiopia completes the eastern trio of sensitive Towards threatened. assemblages: nine of 12 endemics are central Africa, species- and endemic-rich faunas in the 168 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA DIJKSTRA (a)ANDV. CLAUSNITZER(b) (CR) onthelower slopesoftheKenyahighlands. altitudinal occurrence, like(b) altitudinal occurrence, spatialand specieswithrestricted (a), especiallythreatens inD.R.C. ashere foragriculture, The destructionofforest (a) Zambia and northern Botswana whose assessment has benefitted from recent fieldwork. Botswanawhoseassessmenthasbenefittedfromrecent Zambia andnorthern many ofthemhavenotbeenreported since,althoughrecent species havebeendescribedfrom theselocalities,but north-west ZambiabyPinhey(1961b;1984).Numerous (1975); (4)south-westUgandabyPinhey(1961a);and(5) Cameroon byVick (1999);(3)north-eastGabonbyLegrand studied: (1)MountNimbabyLegrand(2003);(2)south-west highlights areas thathavebeendisproportionately well The distributionofDataDefi threatened, orNearThreaten extending from EurasiawhoseAfricanpopulations are (b) (a) (b) (a) Anax bangweuluensis (NT)and cient (DD)species(Figure 5.24b) Platycypha amboniensis ed, with extinction. ed, withextinction.

(b) Ictinogomphus dundoensis

(LC) are twospeciesknownprincipallyfromtheswampsof (LC)are Agriocnemis palaeforma overuse, whichmight endangerapapyrus-endemic suchas poorly protected, thusbeingvulnerable toclearanceand (e.g., Dijkstra2009).Wetlands are alsoeconomicallyvitalbut are importantformany savannahandwoodlandodonates gracile exception of be directly threatened bytheseactivities,withtheprobable and agriculture isdisproportionately high.Few speciesmay surroundings, humanpressure onriparianforests forwood endemics. Becauseoftheircomparativelydryandfi Neodythemis nyungwe gilliesi amboniensis (VU) inCameroon’s BamendaHighlands, populated areas. Examplesare that are fragmentarybynature andlieinfertile anddensely speciesofsubmontaneforestsThis especiallyconcerns main threat toOdonatainAfricaandthetropics worldwide. water bodiesbyerosion, eutrophication andsiltation,isthe agricultural encroachment, andthesubsequentalterationof landscape through deforestation, urbanisationand As humanpopulationsgrow, transformationofthenatural 5.4.2.1 Habitatdegradationanddeforestation 5.4.2 Majorthreats material obtainedthere (Dijkstra this mayrefer tothecollectionheldatthistownratherthan several specieswere describedfrom Dundo(=Chitato),but Suhling further demonstratehowdatadefi Kipping 2000;2003a,b;unpubl.;SuhlingandMartens2007) east NamibiaandadjacentZimbabwe(Pinhey1976;1984; brydeni bangweuluensis in 2010)haspartiallyremedied this.Newrecords of since 2008)andnorth-westZambia(KippingGünther Schütte in2008),south-eastGabon(Mézière andDijkstra fi eldwork insouth-westCameroon (Dijkstra,Kippingand (CR)inTanzania’s eastUsambara mountains, (CR) in western Africa,butshadedrefuges (CR)inwestern alongrivers from anothersucharea –northBotswana,north- et al. (CR)incentralKenya, 2009a). North-east Angola stands out because 2009a).North-eastAngolastandsoutbecause Ceriagrion citrinum , Ictinogomphus dundoensis

© J.KIPPING (NT) in Rwanda, and all Ethiopian (NT)inRwanda,andallEthiopian (NT). et al. Chlorocypha centripunctata ciency can be reduced (cf. ciency canbereduced (cf. (VU)and 2011). Amanipodagrion Amanipodagrion Zygonychidium Zygonychidium and Platycypha Platycypha re-prone Trithemis Anax Anax

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 169 (CR) from Tetrathemis Tetrathemis pits under a diamond © K.-D.B. DIJKSTRA Oreocnemis phoenix Oreocnemis © K.-D.B. DIJKSTRA (a) AND © K.-D.B. DIJKSTRA (a) AND

cial and detrimental (Dijkstra (LC). lter rainwater, and to buffer for spates and to buffer lter rainwater, ies. This suggests that the water quality was ies. This suggests that the water quality ow of clean water is determined by the capacity were found at abandoned were cient despite the extensively damaged surroundings. (a) (b) Mining, such as for gold on the fringes of Ghana’s Atewa Forest Mining, such as for gold on the fringes of Ghana’s Reserve (a), may create habitat for well-dispersing dragonfly species of oen habitats, but may also potentially wipe out endangered species, such as (b) and droughts. as observed in western Small-scale mining in rainforests, Liberia, can be both benefi new scarce pits can create 2007e). Partly overgrown of the localised individuals habitat: emerging godiardi which the Densu supplies one-third of Accra’s water. Torrential Torrential water. of Accra’s which the Densu supplies one-third were levels, and while trees downpours did not alter stream hosted typical along the lower Densu, it still removed largely dragonfl forest suffi The steady fl on the plateaus and in theof the soil, swamps and forest and fi valleys to store the bauxite-rich Mt. Mulanje in Malawi. Proischnura rotundipennis Proischnura

ow uctuations appearing Proischnura and 1996), and the Elattoneura pluotae Oreocnemis phoenix et al. (NT). Not all damming Paragomphus cataractae ood critical riverine habitat,ood critical Agriocnemis pinheyi ood habitat of been inundatedy habitat have already tting from the constructions of trout tting from Africallagma sapphirinum Mesocnemis dupuyi ) benefi ned to rapids and waterfalls of large rivers, large ned to rapids and waterfalls of (a) (b) E.R. NIELSEN (b) downstream for downstream to all species, with some endemics in harmful events are South Africa (e.g., rotundipennis dams (Samways 1989; Samways following dam construction. For example, Lake Volta in Ghana Volta following dam construction. For example, Lake in reservoirs the largest and Lake Kariba on the Zambezi are (over 8,500km²) and volume (up tothe world by surface area the Akosombo before No data exist from 185km³), respectively. completed in 1965 and 1958, butand Kariba Dams were in Ghana Volta O’Neill and Paulson (2001) surveyed the Black surveyed theabove the new Bui Dam and Dijkstra (2009) power above the envisaged Igamba Malagarasi in Tanzania plant. Even smaller dams can fl such as rapids and gallery forests. 5.4.2.2 Dams of dragonfl Huge areas (NT) is confi electric power plants planned at from coming under threat and at PopaEpupa Falls on the Kunene (cancelled in 2008) fl downstream Falls on the Okavango. Dams also impact and sedimentation patterns. in the upperregimes Damming Gambia catchment may fl erratic water level fl and create (CR) upstream southern African endemic soon after a pond was created for plant conservation (Steytlersoon after a pond was created and Samways 1995). 5.4.2.3 Mining wealth of mineral for its disproportionate Africa is renowned often extracted by open-pit mining, These are resources. leading to extensive damage of the landscape. This is composed entirely in highlands that are especially problematic ore of economically valuable deposits, examples being iron in Mount Nimba (see 5.3.2.2) and bauxite in Mulanje (5.3.3.4). Exploitation of the latter could wipe out (CR) completely. Dijkstra (2007f) observed the importance (CR) completely. Atewa catchments in Ghana’s of highlands as freshwater major rivers in this Range, which has a bauxite cap. Three of have their headwaters there, densely populated region While large dams like Akosombo in Ghana (a) can flood greatWhile large dams like Akosombo odonate habitats, small dams can create expanses of varied like the South African endemic (b) valuable habitat for specialists 170 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA all localised endemics(Samways andTaylor 2004;Samways natural habitats and affecting manywidespread speciesand vegetation along streams inSouthAfrica, radicallyaltering Australian wattles( Similarly, invasivealientrees canbeakeythreat: riparian (Shelton, Samwaysunpubl.). (see 5.3.3.2),andresearch ontheirimpactisunderway likely toendangersensitivespeciesliketheCapeendemics headwaters (orthosewithindigenousspecialisedfi In SouthAfrica,introducing trout toformerly fi sp.) introduced tocombatmalariavectors(Dumont1991). extirpated insouth-westLibyabymosquitofi such effects inAfrica,but trutta rainbow trout ( (regionally) alienspecieslikeNileperch ( introduction tostreams andwetlands, particularlyof Fish are importantpredators ofodonatelarvae.Their 5.4.2.4 Alienplantsandfi comparison toapristinestream. was suitablefor of drainagestreams. OnesuchstreamLiberia inwestern pits alsoaffects thefl Reduced motionandincreased sunningofwaterinopen reduces visibilityforlarvae,thelatteraffects thesubstrate. turbidity, andprobably siltation,ofstreams. Theformer Moreover, thedrainageoftheseminesleadstoincreased dispersing speciesthatoccurthroughout Africa(see5.3.1). closed canopy. Openpits,however, are colonisedbywell- (b) (a) ), canhaveasevere impact.There are fewstudiesof Oncorhynchus mykiss S. fumosa Acacia ow, oxygenandtemperature regimes Agriocnemis sania sh sp.)mayovergrow thenatural (NT),butheldlownumbersin © at theBlydeRiver(b). Pseudagrion newtoni like(a) been removed, invasive alienplantshave known fromsiteswhere only Africa endemicsare South Some threatened W. TARBOTON (b)

) orbrown trout ( E.R. NIELSEN (a) AND E.R. NIELSEN(a)AND Lates niloticus mayhavebeen sh ( sh Gambusia sh-free Salmo Salmo s) is sh) (VU) (VU) ), ),

Areas withalmost norecords butalso very fewexpected example, inthe EthiopianandeastZimbabwe Highlands. confi France andItalycombined.Surveys are alsoneededto west Sudan(Figure 5.2) isalmostthesizeofBelgium, includes muchoftheCentralAfrican Republicandsouth- an ideaofscale,the1200x800km “emptyquarter”that Angola.Toof CongoandcentralD.R.C.to northern give Nigeria, throughCameroon,Republic eastern almost uninterrupted,understudiedswathefrom southern Figure 5.3).CentralAfricastandsoutinparticular, withan most areas despitearecent surge inrecords (see5.1.2.2; The mainprioritynowisfi 5.4.3.1 Fieldsurvey 5.4.3 Research action become NT)onSocotra. African,and (NT) innorthern Paragomphus sinaiticus in thisregard are et al. competition betweenspecies(cf.Martens well asindirectof onesbyalteringwell-establishedpatterns may havedirect effects bydestroying larvalhabitats,as to intermittent,ephemeralorevenuninhabitableones.This Africa,transformingraresouth-western perennial sources pollution are mostcriticalindryenvironmentsand innorthern mortality (Schuran,Kippingunpubl.).Water abstractionand semi-artifi Botswana,testedunder the OkavangoDeltaofnorthern but aerialsprayingofDeltamethrintocontrol tsetsefl Little isknownabouttheeffects ofpollutantsonOdonata, increasing useofpesticidesagainsthumandiseasevectors. development andutilisationofwaterresources, there isan change scenarios.Intandemwithpollutionarisingthrough increasing problem, particularlyconsideringglobalclimate irrigation andindustryisatpresent alocalbutpotentially 2003). Similarly, waterabstractionforhumanconsumption, because manyOdonataare notsalt-tolerant(Suhling regions, asreported inSouthAfrica,mayhaveastrong effect increase. Riversalinationduetointensiveagriculture indry Africa’s industrialandagriculturalsectorstheseimpactswill a localproblem atpresent. However, withthedevelopmentof developed. Chemicalandorganic pollutionistherefore atmost while industryandchemical-intensiveagriculture are under- Human populationsinmanypartsofAfricaare still low, 5.4.2.5 Water extractionandpollution level ofthreat theypose(Samways sites where alientrees havebeenremoved, illustratingthe and Metacnemis angusta endemics (SamwaysandSharratt2010).Some,suchas species (MagobaandSamways2010)threatened these aliensresults inarapidrecovery ofbothwidespread 2006b; SamwaysandGrant2006b).However, removal of rm thestatus of(probably) threatened species,for Pseudagrion newtoni 2006). Examples of species of global or African concern 2006).ExamplesofspeciesglobalorAfricanconcern cial conditions,affects bothimaginalandlarval Calopteryx exul Calopteryx (VU), s (EN eldwork, whichisstillrequired in (VU)are indeedonlyknownfrom Proischnura polychromatica RG ) and (EN), Azuragrion granti et al. Cordulegaster princeps Gomphus lucasii 2005). 2005). et al. 2010; Suhling 2010;Suhling (LC; will (LC;will ies in in ies (VU), (VU), et al. (CR) (CR)

CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 171 ies are well ies are Azuragrion ies and damselfl (LC) from Socotra. © J.-P. BOUDOT (a) AND R. KETELAAR (b) © J.-P. Natural aquatic habitats and their inhabitants are under pressure by increasing human demand for water especially in arid environments, as can be imagined on the Oued Chouf in Morocco (a) and for an endemic of a dry island, like (b) granti Southern Mozambique; and Eastern Angola and western Zambia. Southern Tanzania and northern Mozambique; Southern Tanzania ciently investigated to determine reliable trends. To To trends. ciently investigated to determine reliable cient is the knowledge of population ecology. As ecology. cient is the knowledge of population (10) (11) (12) 5.4.3.2 Ecological research Although, in general, dragonfl known ecologically (Corbet 1999), knowledge of tropical Especially and particularly African species is often poor. defi Red List Criteria A, C and D could the IUCN a result, be applied in only a few cases for the African Odonata could be population trends assessments. Even where for instance by indirectly, inferred estimated, these were as the habitat of rates of habitat loss. This is problematic, known, being based mostly most species is not precisely on sketchy descriptions of observation localities. At most and Cape endemics may have been a handful of Maghreb suffi population assessments in the future, reliable enable more least) of (at habitat requirements and detailed trends selected species should be investigated. Candidates are has been inferred species for which population decrease because this may allow for habitat destruction, from This habitat and population trends. between correlations monitoring of population size at selected regular requires sites, carried out by local researchers. North-west Upper Guinea: especially the Fouta Djalon in Guinea; River North-west Lower Guinea: especially the Cross State and Niger Delta in Nigeria; south from area South-east Lower Guinea: most forest Angola; to north-west Cameroon “Cuvette” and south Congo Basin: especially the West Salonga National Park); (e.g., along the Likouala River, Albertine Slope and Highlands in eastern D.R.C. West Maiko and Kahuzi-Biega National (e.g., Ituri Forest, Parks); and Guinea, southern Mali and northern Côte d’Ivoire; Northern northern Nigeria, through Cameroon, southern Chad and the Central African Republic to southern Sudan;

Angolan Escarpment and Highlands. Northern and eastern Tanzania; (b) (a) species have the lowest research priority, for example, east priority, species have the lowest research Kenya and south-west Botswana. north-west to south-east), priorities (ordered The greatest and threatened with the most likely discovery of new species are: (1) (2) (3) (4) (5) (6) with open medium-rich regions Of secondary priority are especially: habitats or fragmented (highland) forests, (7) (8) (9) 172 CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA preferences, are Chlorocyphidae, overall diversityinspeciesnumbersandecological scientifi ecology, Odonataare themostsuitablegroup tofi reasonable knowledgeoftheirtaxonomy, distributionand to focusonaquaticgroups equippedforfl of phylogeneticstudiesonAfricananimalsare estimated relationships withhostbiota.Nonetheless,fewerthan2% by humandisturbance,andinmanyinsectsintricate by theirconfi areFor example,patterns complicatedinfi geological, ecologicalandhumanfactorsare limited. fl by fl can potentiallyalsorespond instantlythrough dispersal such asOdonataare affected directly bychangeand and vegetationofaquatichabitats.Amphibiousinsects Climate changeaffects thedensity, size,permanence,fl from fi combining ecological,geographicandphylogeneticdata 5.2.2.1; 5.2.2.2).Diversifi history canaidtheconservationofpresent diversity(see An understandingofenvironmental andevolutionary 5.4.3.3 Understandingchange ferox an excellentexamplebeingthestudyof Malawi. Alsomore autecologicalstudieswouldbeuseful, at monthlyintervals,ascarriedoutbyParr(1984)in is possiblethrough regular countsofadults,forexample, from whichmuchlifecycleinformationcanbeinferred. This droughts. Afi if climatechangeleadstomore regular orsevere seasonal most chlorocyphids andcalopterygids,maybeimpacted and Lindeboom2002)forest stream species,suchas tree-hole breeders like explored tounderstandtheirsensitivity. Forexample, The lifecyclesofforest speciesinparticularmust to be 1998; Samraoui2009;Suhling fringes(e.g.,Samraoui andsouthern arid northern the possibleexceptionofsomespeciesoncontinent’s Life cyclesofAfricanOdonataare alsopoorlyknown, with species, where are sensitiveor tolerant ones? Where example, where (regions, habitats) are olderor younger viewed together tohelpusanswerfurtherquestions –for similar events?Thirdly, ecologyandevolutioncanbe ages, andwhethergroups showsimilarresponses to to helpanswerquestionsabout whatare species’relative disturbance orchange.Secondly, historicdepthisadded can thenbetterunderstandhow theymayrespond to factors, andwhatare theirdispersalcapacities?We tolerant, are speciesinrelation towhich environmental (see also5.4.3.2)toquestionssuchashowsensitive,or This research wouldfi Trithemis 5.14), iers” are honestindicatorsalsobecausecomplicating ight (“rangefollowschange”).Such“freshwater byMiller(1964). eld surveys,GISanalysisandmolecularresearch. Elattoneura c gap.Suitableradiationsforstudy, representing (Damm nement toriverbasins,inmanyvertebrates rst stepistoobtainmore phenologicaldata, et al. , Pseudagrion rstly provide basicecologicalanswers 2010). Coryphagrion grandis Coryphagrion cations canbeunravelledby (Dijkstra et al. Chlorocnemis 2003,2004,2005). et al. ight. Givenour Ictinogomphus (Clausnitzer sh patterns patterns sh 2007b)and (Figure l this ll et al. ow

(5) (4) (3) (2) (1) conservation inAfricaas: to promote dragonﬂ poor. Recentdevelopmentsmakeittimelyandfeasible has increased, accesstorelevant knowledgeremains Although attentionforaquaticbiodiversityconservation of climatechange,toolstoassessimpactsare needed. ecosystems, nowexacerbatedbythehydrological impact As mankindcontinuestodevastatefreshwater 5.4.3.4 Informationdisclosure diversity possible? close relatives overlappingextensively, howishighlocal If radiationsare explosiveandspeciesadaptable, with or recent? Whatistheinﬂ assemblages inforests orstreams relatively sensitiveand/ ones donotsurvivelong?Forinstance,are species older specieshavewidertolerances,becausesensitive differ foraquatic(larval)andterrestrial (adult)habitats?Do savannah), andwhichshiftsare prevalent? Dotheserates ancestral, howfrequent are ecologicalshifts(e.g.,forest to by immigrationorlocalspeciation?Whichecologiesare might speciesoriginateorsurvive?Isassemblagechange

and Tarboton 2002;2005;Dijkstra andLewington worldwide boom inpopularliterature (e.g.Tarboton 2001; 2005;Niba andSamways2006),asseen inthe gained broad public appeal (e.g.,SuhandSamways Being colourfulandconspicuous, dragonfl Africa’s freshwater; and second-largest rainforest, containing one-quarterof environmental health of theCongoBasin,world’s Degradation (REDD)incentralAfricadependsonthe Dam, andReducedEmissionsfrom Deforestation and generating twicethepowerofChina’s Three Gorges For example,thefeasibilityofGrandIngascheme, water andenergy are becomingprominent resources. deposits are beingincreasingly exploited, andalso (e.g., Dijkstra2007b,e,f;2009).Africa’s richmineral are vulnerable,forexample,bymininganddamming impact assessments,especiallywhere aquaticsystems Odonata are increasingly employedinenvironmental 2008); measuring habitatrecovery (SimaikaandSamways method (SimaikaandSamways2009b)usedin 2010), andsubsequentlyappliedasasite-selection tool forhabitatintegrity(SimaikaandSamways2009a; South Africaasarapidenvironmental assessment The Dragonfl report; database forallAfricanspecies,assummarisedinthis study) resulted inRedListassessmentsandarecord The IUCNFreshwater BiodiversityAssessment(current Dijkstra andClausnitzerinprep.; Samways2009); and Tarboton 2002;2005;SuhlingandMartens2007; nearly 75%ofspecies(becoming)available(Tarboton revised (Dijkstra2007d),withidentifi The taxonomyoftheover700specieshasbeenlargely y BioticIndex(DBI)wasdevelopedin ies asafocalgroup infreshwater uence ofecologicalinteractions? cation literature for is have ies CHAPTER 5 | DRAGONFLIES AND DAMSELFLIES OF AFRICA 173 uctuations is © K.-D.B. DIJKSTRA ies, require shade in their adult stages. ies, require sh should generally not be released, and sh should generally not be released, ow are needed, the simplest being minimising needed, the ow are uctuations may also seriously impact aquatic life uctuations may also seriously impact ood plains. ow by recycling water and the use of settling ponds. ow by recycling In large-scale landscape altering projects like open-cast landscape altering projects In large-scale mining and plantations, damage to the watershed can zones of natural buffer be minimised by leaving broad water bodies (rivers, inundation zones, vegetation around removal swamps) (Kinvig and Samways 2000). Forest organisms, should generally be avoided, as many freshwater including dragonfl be guaranteed to avoid spates and droughts. The impact be guaranteed to avoid spates and droughts. systems is obvious, but irregular on freshwater of drought water fl of the micro-climate cycles, for example, by affecting The and disturbing adult emergence. habitats breeding if a can be reduced impact of dams, at least downstream, with normal seasonal fl natural water regime Samways 1996; Stewart and Samways (Clark and retained 1998). Alien fi banks stream from should be removed invasive alien trees and fl such as in irrigation and water, usage perturbs Where and rivers to avoid siltation of streams mining, measures the outfl from outfl y cation eld photos for (both daily and ow regimes s cation tools and accessibility to cient spill-over and regular discharge must discharge cient spill-over and regular 2006; Suhling and Martens 2007; Samways 2009) and 2006; Suhling and Martens 2007; Samways of the volunteer workforce. growth Furthering their role in freshwater conservation in Africa in freshwater Furthering their role identifi improved requires data on status, ecology and distribution, on which DBI is necessary to assemble, generate based. It is therefore and disclose taxonomic and ecological data, identifi tools, range maps, specimen scans and fi but knowledge all species. Species diversity is greatest, Gabon and the least, in countries such as Cameroon, the Congos (see 5.4.3.1), so the stimulation of dragonfl research in francophone Africa is especially important. research 5.4.4 Conservation action To conserve Odonata, and the freshwater biodiversity Odonata, and the freshwater conserve To it is necessary to maintain the structural they represent, integrity of both larval and adult habitats, i.e., water bodies to avoid landscape. Many measures and their surrounding unnatural fl siltation and erosion, or piping When damming streams simple. seasonal) are springs, suffi Odonates require diverse and structured habitats for their survival. The Malagarasi in western Tanzania shows how the Odonates require diverse and structured habitats for their survival. The Malagarasi in western Tanzania streambed create can different freshwater habitats, like rapids, pools and swamps. morphological diversity of a single river’s Gallery forest provides shelter in the drydry season, which has left the surrounding hills and inhospitable. 174 CHAPTER 5 | SPECIES IN THE SPOTLIGHT Senckenberg MuseumofNatural HistoryGörlitz, PF300154, D-02806Görlitz,Germany ¹ K 8^lXi[`Xef]B\epXËjnXk\ij_\[1DXXk_X`Ëjcfe^c\^ recognised for her effort to advocate recognised forher efforttoadvocate this prize.Wangari Maathai was African womantobehonoured with laureate, Wangari Maathai,thefi in honourofthe2004Nobel Peace Notogomphus fl race” ofanotherclubtailspecies, been misidentifi Museum (boththesespecimenshad in London’sNatural History V.G.L. vanSomeren)turnedup early EastAfricannaturalist Mountains bythepioneering southern slopesoftheAberdare Katamayu Forest (onthelower, collected duringthe1950sin other (male)specimens,both as adistinctspecies)andtwo (hitherto ithadnotbeenrecognised the National Museums ofKenya turned upinthecollectionof between Nakuru and Kericho) (on theEasternMauEscarpment Watulege intheMarioshoniForest one male,collectedin1970byB. and morespecimensdiscovered: the wild,collectionswerebrowsed discovery ofthisnewspeciesin history museums.Following the old Odonatacollectionsinnatural of siftingpainstakinglythrough up, notinthewild,butasresult world inrecentyearshaveturned new speciesidentifi years) wasararethrill.Indeed,most species (thefi studied inKenya;sofi since the1950s,beenfairlywell late discovery. Dragonfl forest streamscanexplainitsrather restricted occurrenceinmontane bright greendragonfl National Park. Itisaverystriking, River, inKenya’sMountElgon The newdragonfl Jg\Z`\jèk_\jgfkc`^_k 2000 alongtheRongai was fi ( longleg he Maathai’s Notogomphus maathaiae rst forKenyainmany rst discoveredin avifrons ed asa“Kenyan y was named y wasnamed ed aroundthe y andonlyits nding anew ). ies have, rst rst ) montane discovery oftheapparentlytruly of manymontaneendemics.The montane faunanorforthepresence are neitherknownforhavingarich the survivalofuniquebiodiversity. benefi and theriversthatfl against poverty. Safeguardingforests, world’s naturalresourcesinthefi importance oftheprotection choice for2004emphasizesthe forests. TheNobel Committee’s last remainingandfastshrinking focusing ontheprotectionofAfrica’s sustainable humandevelopmentby dragonfl surprising. Othernotablemontane streams above2,200m,istherefore to berestrictedmontaneforest slopes of Mt. Kenya and the eastern slopes ofMt.Kenya andtheeastern and isendemic to thewestern P. amboniensis Tanzania between1,500and3,000m. forest streamsinKenyaandnorth are largelyconfi and amboniensis The Maathai’s longleg ( The easternAfricandragonfl Atoconeura kenya ts mankind and also ensures ts mankindandalsoensures ies inKenyaare N. maathaiae , Pseudagrion bicoerulans

occurs below 2,000m occurs below2,000m ned to highland ned tohighland ow from them, ow fromthem, . All three . Allthree , which seems , whichseems Notogomphus maathaiae Platycypha Platycypha ies ies ght ght the watershed”. as thisonearedubbed “guardiansof source. To thisend,dragonfl and acleandependablewater foothills, byguaranteeingstable soils dragonfl riverside forestsnotonlyhelpsthis watershed protection.Conserving and asafl as anindicatorofhealthystreams Endangered (CR). It therefore serves Endangered (CR).Itthereforeserves rare dragonfl densely populatedhighlands,this As deforestationcontinuesinthe have sufferedsignifi recent decades,andthespeciesmust have beenwidelydestroyedin high mountainforests.Suchhabitats known fromclearstreamsinthree List ofThreatenedSpecies.Itisonly Endangered (EN)onTheIUCNRed found below2,000m. of altitudesabove2,500mandisnot on theotherhandischaracteristic the mountainslopes. agriculture anddeforestationon particularly vulnerabletoexpanding slopes oftheAberdares,makingit The Maathai’s longleg is assessed as longlegisassessedas The Maathai’s ) (EN). y, butalsothepeopleof agship species for urgent agship speciesforurgent y maybecomeCritically

VIOLA CLAUSNITZER cant declines. Clausnitzer, V¹ P. bicoerulans ies such CHAPTER 5 |

SPECIES IN THE SPOTLIGHT

The globe skimmer (Pantala ﬂ avescens) (LC). © VIOLA CLAUSNITZER

Jg\Z`\j`ek_\jgfkc`^_k K_\^cfY\jb`dd\i1feXaflie\pXifle[k_\nfic[

Clausnitzer, V¹

he globe skimmer, Pantala migration, dying on the way and fl avescens, is arguably the leaving its offspring to complete the most successful dragonfl y journey. This in itself is remarkable, in the world, with a but the migration distance covered is Kdistribution range extending to all truly impressive, and makes the globe continents except Antarctica. In skimmer the strongest migrating tropical areas and in North America insect species known. its numbers often build into large The globe skimmer breeds aggregations which become true in small, bare, stagnant water migratory swarms. In Africa large bodies, and is commonly found in swarms of the species, which is at an altitude of 1,000m, where it temporary rain pools. As a strong known locally as the “helicopter unwittingly provides an important migrant it is specialised to colonise insect”, are taken by the people to food source for migrating birds these temporary habitats as its larvae indicate to coming of the rains. It taking the same route. It is thought have evolved to complete their life- is also known as the “wandering that P. fl avescens takes advantage cycle in one month, which is shorter glider”, as its wings are large of the moving weather systems and than most species. The advantage and broad, helping it to glide for monsoon winds to complete a truly of colonising temporary rain pools, exceptionally long distances. epic migration from southern India and of a short development time, is Although the migratory swarms to east and southern Africa, and back reduced competition and predation, of this species are well known, again. This represents a round trip of as few other species are able to it was only recently discovered 14,000 to 18,000km that takes several colonise these pools so quickly. that it regularly migrates over generations to complete. How each The success of this adaptation incredible distances. For example, dragonfl y knows the migration is evidenced by the unusually the species has been recorded as route remains a mystery, given that high numbers in which the globe travelling thousands of kilometres no individual completes the full skimmer occurs.

¹ Senckenberg Museum of Natural History Görlitz, PF 300154, D-02806 Görlitz, Germany 175 176 CHAPTER 5 | SPECIES IN THE SPOTLIGHT 1,000km away assume, butinMadagascar, African mainlandasonemight Senckenberg MuseumofNatural HistoryGörlitz, PF300154, D-02806Görlitz,Germany ¹ K [Xdj\cÕ K_\G\dYX]\Xk_\ic\^1X]iX^`c\ of material lyinginthestream. laying) takesplaceondeadplant carefully, andoviposition (egg to arrive.Theyapproachfemales water whilewaitingforfemales to, thesluggishstream,facing on twigsandleavesabove,orclose and denserainforest.Malessettle the darkunderstoryofmoist of around1,500ha. status in1959,withagazettedarea Forest, attainedForest Reserve remaining forestpatch,theNgezi kilometres oforiginalforest.This 1830, leavingonlyafewsquare (cloves, cardamom)beganin clearance forcashcropplantations covered allofPemba Island.Forest a tropicalmoistforestthatonce the onlyremaininglargepatchof coast ofTanzania. Ngezi Forest is offshore Islandonthenorthern Forest onPemba Island,asmall along theonlystreaminNgezi delicate damselﬂ formally describedin2007.This only discoveredin2001andwas pembipes Remarkably, thenearestrelatives The Pemba featherlegdwellsin Platycnemis pembipes Jg\Z`\j`ek_\jgfkc`^_k quill-like shins, (CR), namedforitswhite ( he Pemba featherleg Platycnemis pembipes Remarkably, thenearest relatives of y wasdiscovered are found not on the are found notonthe arefound

was p`eX]iX^`c\i\Xcd ) Platycnemis Platycnemis pembipes 1,000km away. Theoriginof might assume,butinMadagascar, not ontheAfricanmainlandasone island’s species arealsofound Pemban taxa. While manyofthe P. pembipes The Madagascanconnectionof carried thesedamselﬂ to Septembercouldpossiblyhave coast north-westwardsfromJune which followstheEastAfrican Madagascar. Thestrongmonsoon the MozambiqueChannelfrom through wind-aideddispersalacross The Pembafeatherleg( on Pemba has parallelsinother

is mostlikely ies toPemba. Platycnemis pembipes P. P. the brinkofextinction. which arebringing thespeciesto pollution andforestclearance, disturbance suchasstream threats, includinghabitat restricted rangeandongoing Species becauseofitsvery IUCN RedListofThreatened as CriticallyEndangeredonThe coastal forestsonthemainland. assemblage ofspeciesalsofoundin odonates representatypical In otherrespects,Ngezi Forest elements foundincombination. Indian speciesandAfromontane forests, Madagascanlinks,eastern restricted tocoastalEastAfrican composition, withtreespecies Ngezi ispeculiarin itsﬂ For instance,themoistforestof unique combinationofspecies. mainland, recognizedforits 1,000km²) andproximitytothe is, despiteitssmallsize(about in continentalAfrica,Pemba Platycnemis pembipes ) (CR). ©

VIOLA CLAUSNITZER Clausnitzer, V¹ is assessed oristic CHAPTER 5 Jg\Z`\j`ek_\jgfkc`^_k

8nfe[\i]lcnXk\ii\c`Z | n`k_XeXeZ`\ekc`e\X^\ SPECIES IN THE SPOTLIGHT Samways, M¹

he gilded presba, water courses of the Cape. They The cherry on the icing was Syncordulia legator (VU), shade out the native habitat and provided by geneticist Jessica Ware, is not only a stunning render conditions unsuitable for who examined the DNA of the insect but one with great many sun-loving Cape endemic genus Syncordulia, and discovered Kcharisma. Until recently nothing dragonfl y species. However, at this another new species, the rustic was known of the species, with site, where the gilded presba was presba (Syncordulia serendipator). only a couple of broken specimens fi rst seen alive, alien trees had been Jessica found that the species in in museums hinting at its existence. removed as part of the national the genus diverged some 60 million Then one day in October 2006, Working for Water Programme. years ago. Although these species sitting on a bush in one of the This was originally proposed to had survived the various comings most beautiful settings of the improve hydrology and give jobs and goings of cold and warm eras, ancient Cape Fold Mountains, this but, as it turned out, was also in the geological blink of an eye stunning dragonfl y was found, amazingly encouraging for the they were almost rendered extinct freshly emerged, and glistening in return of this and other threatened by invasive alien trees. They the early morning sun. This was freshwater species. have been given a reprieve by the not a small insect, and was even removal of alien trees through the declared as being the size of a Working for Water Programme “young elephant”, which although in South Africa. This reprieve an exaggeration, nevertheless came about only because the key emphasizes how extraordinary it is threat was identifi ed (alien trees) that such a large and magnifi cent and dealt with, allowing narrow insect could have been overlooked range endemics, like the gilded in the past. presba, to return. This emphasizes There are two reasons why this that ancient endemics can be species had not been recognized surprisingly resilient when they are earlier. The fi rst relates to its given the opportunity by removing biology. It emerges early in the major human-induced threats. summer season, and immediately Without habitat restoration such departs from its natal river and fl ies as this, it is quite possible that high into the rugged mountains, the gilded presba would have where it cruises swiftly over the The gilded presba (Syncordulia gone extinct without ever being bushy fynbos vegetation, unique to legator) (VU). © MICHAEL SAMWAYS discovered. the Cape Floristic Region in South Africa. Although a week after the fi rst sighting a visit to the site produced some females, equally Although these species had beautiful with golden wings, a few survived the various comings weeks later none were to be seen – all having departed for the upper and goings of cold and warm slopes. The second reason why this species had not become known was eras, in the geological blink of an that its habitat had been threatened eye they were almost rendered by the impact of invasive alien trees. The trees, mostly Acacia and extinct by invasive alien trees native to Australia, thrive along the

¹ Department of Conservation Ecology and Entomology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa 177 Chapter 6. Freshwater crabs of Africa: diversity, distribution, and conservation.

Cumberlidge, N.¹

¹ Department of Biology, Northern Michigan University, Marquette, Michigan 49855, USA

The Purple March Crab Afrithelphusa monodosa (Endangered) which lives in swamps and year-round wetland habitats in north-western Guinea where it is known from only a few specimens from two localities. This species is clearly a competent air-breather and has a pair of well-developed pseudolungs. It is mainly threatened by habitat loss and degradation. © PIOTR NASKREKI

An unidentiﬁ ed freshwater crab species within the family Potamonautes. This specimen was collected in central Africa, a region noted for its limited ﬁ eld sampling. © DENIS TWEDDLE The Purple March Crab Afrithelphusa monodosa (Endangered) which lives in swamps and year-round wetland habitats in north-western Guinea. © PIOTR NASKREKI

Potamonautes lirrangensis (Least Concern), a relatively abundant and widespread species found in large slow ﬂ owing rivers in rainforests across central and eastern Africa. © DENIS TWEDDLE

CONTENTS 6.1 Overview of the African freshwater crab fauna 180 6.1.1 Biogeographic patterns 182 6.2 Conservation status 183 6.3 Patterns of species richness 184 6.3.1 All freshwater crab species: interpretation of distribution patterns 186 6.3.2 Threatened species 187 6.3.3 Restricted range species 188 6.3.4 Data Defi cient species 190 6.4 Major threats 191 6.4.1 Habitat destruction 191 6.4.2 Pollution 192 6.4.3 Natural predators and competition with introduced species 192 6.4.4 Taxonomic issues 193 6.5 Research actions required 193 6.6 Conservation recommendations 195 Species in the spotlight – A unique species fl ock of freshwater crabs in Lake Tanganyika: a model for studies in evolution and extinction 196 Species in the spotlight – The East African tree hole crab – spectacular adaptation 198 180 CHAPTER 6 | FRESHWATER CRABS crab insmall streamsthroughout therainforest andsavanna zonesofwestern Africa. 2006; Cumberlidge Potamonautidae (Cumberlidge1999,2009a,b;Daniels of freshwater crabsin10generathatallbelongtothe Afrotropical zone.ThispartofAfricaincludes119species (Socotra, theSeychelles,andMadagascar)thatalsoliein IndianOcean Desert, butexcludestheislandsofwestern here includesall43AfricancountriessouthoftheSahara et al 1896 (Yeo Potamonautidae Bott,1970,andPotamidaeOrtmann, The CommonCreek Crab genera ( where there are three speciesoffreshwater crabsintwo freshwater crabsare Morocco, Algeria,Tunisia, andEgypt, Afrotropical Africathat host zone.Thecountriesofnorthern zone, andtherest oftheAfricancontinentthatlies inthe AfricainthePalaearcticincludes northern zoogeographical focuses onthefreshwater crabsofcontinentalAfrica, which 1955; Cumberlidge1999;Yeo and smallerwaterbodies(Rathbun1921;Balss1936;Bott habitats, from mountainstreams tolarge lowland rivers conspicuous crustaceansare present inalmostallfreshwater inhabiting Africaninlandwaters,andtheselarge and Freshwater crabsare amongthemostimportantinvertebrates 6.1 and Ng2009;Cumberlidge . 2009).ThepartoftheAfrotropical zonedealtwith crab fauna Overview oftheAfricanfreshwater Potamonautes et al . 2008; Cumberlidge 2009a,b; Cumberlidge . 2008;Cumberlidge2009a,b; et al and . 2008;Yeo Liberonautes latidactylus et al etal Potamon . 2009). . 2008). The present work . 2008).Thepresent work et al ) and two families, ) andtwofamilies, . 2008; Cumberlidge . 2008;Cumberlidge (Least Concern)is themostcommonandfrequently caughtfreshwater et al . Cumberlidge family, thePotamonautidae(Bott1955;Cumberlidge1999; and specieslevelswith100%endemicityforthepredominant freshwater crabfaunaishighlyendemicatthefamily, genus, al (154 genera,849species,twofamilies)regions (Yeo genera, 298species,twofamilies)andOriental/Palaearctic seen inthefreshwater crabfaunasoftheNeotropical (50 120 species,twofamilies)(Table 6.1)ismarkedlylowerthan The totaltaxonomicdiversityofcontinentalAfrica(11genera, et al Tavares 2006;CumberlidgeandDobson2008; 2004; CumberlidgeandReed2004,2006; et al 1998, 1999,2009a,b;CumberlidgeandBoyko2000;Corace less diverseregional fauna(Bott1955;Cumberlidge1997, Africa(fourspecies,twogenera)eachhavea and northern Africa(20species,onegenus) six genera),whilesouthern (33 species,threeAfrica(24 genera),andwestern are Africa(44species,three eastern genera),central Africa more thanoneregion. Forexample,themostdiverseregions composition, withonlyafewspecies(14outof120)foundin and eachofthesehasadistinctfreshwater crab faunal purposes ofthisproject, subdividedintothefi . 2008;Cumberlidge . 2001;Cumberlidge . 2009) . 2009) et al © PIOTRNASKREKI . 2009). The African continent is, for the . 2009).TheAfricancontinentis,forthe et al et al . 2002;CumberlidgeandVanini . 2009).ContinentalAfrica’s ve main regions, ve mainregions, et et CHAPTER 6 | FRESHWATER CRABS 181 Potamonautidae, Potamonautinae Potamonautidae, Hydrothelphusinae Potamonautidae, Hydrothelphusinae Potamonautidae, Hydrothelphusinae Potamonautidae, Hydrothelphusinae Potamonautidae, Hydrothelphusinae Potamonautidae, Hydrothelphusinae E Potamonautidae, Potamonautinae E Potamonautidae, Potamonautinae E Potamonautidae, Potamonautinae N Potamidae, Potaminae W Potamonautidae, Potamonautinae W Potamonautidae, Potamonautinae W Potamonautidae, Potamonautinae 1 1 1 1 2 1 2 1 1 1 1 1 1 1 No. Genera Region subfamily Family, N = northern W = western Africa, E Africa, C = central Africa, 1 2 1 1 2 1 2 No. Species Country 56 Nigeria Liberia7 Guinea89 Côte d’Ivoire10 Ghana11 Togo Sierra Leone1213 Benin 1014 Chad 8 15 Mali16 Mauritania 5 17 3 6 Burkina Faso18 Niger19 D. R. Congo 6 2 20 Cameroon W 4 321 Congo22 Potamonautinae Potamonautidae, Gabon23 3 2 Central African Republic24 W 2 Equatorial Guinea/Bioko 225 2 Angola (Cabinda) 1 2 Potamonautidae, Potamonautinae 26 Sudan W 2 Somalia 24 W27 1 Potamonautidae, Potamonautinae 1 428 1 13 1 Ethiopia 2 Potamonautidae, Potamonautinae 3 Kenya W29 11 3 W30 8 1 Uganda Potamonautidae, Potamonautinae W 2 4 W 131 W Rwanda Potamonautidae, Potamonautinae 132 Potamonautidae, Potamonautinae Burundi Potamonautinae Potamonautidae, C 3 Potamonautidae, Potamonautinae Tanzania W33 2 C C 1 3 Potamonautinae Potamonautidae, 34 C 1 Malawi Potamonautidae, Potamonautinae 35 Potamonautidae, Potamonautinae Potamonautinae Potamonautidae, C Angola36 Potamonautidae, Potamonautinae 6 Botswana C C37 Potamonautidae, Potamonautinae 2 13 Lesotho38 Mozambique Potamonautidae, Potamonautinae 12 Potamonautidae, Potamonautinae 39 Namibia 1 440 South Africa E41 1 Swaziland 24 42 2 Zambia Potamonautidae, Potamonautinae E Zimbabwe 1 4 1 Potamonautinae Potamonautidae, 5 2 E 3 1 E Potamonautidae, Potamonautinae 1 E 14 1 4 Potamonautinae Potamonautidae, 1 Potamonautinae Potamonautidae, 1 1 1 E 3 1 S 3 S 1 Potamonautidae, Potamonautinae S S 1 Potamonautidae, Potamonautinae Potamonautidae, Potamonautinae S Potamonautidae, Potamonautinae 2 Potamonautinae Potamonautidae, 1 S S Potamonautinae Potamonautidae, Potamonautinae Potamonautidae, S Potamonautidae, Potamonautinae S Potamonautidae, Potamonautinae Potamonautinae Potamonautidae, 12 Morocco3 Algeria4 Tunisia Egypt 1 1 1 1 1 1 1 N N Potaminae Potamidae, N Potamidae, Potaminae Potamidae, Potaminae Table 6.1. Numbers of species, genera, families and subfamilies, and their regions of occurrence, for each of for each of occurrence, regions and their and subfamilies, families genera, of species, 6.1. Numbers Table crabs. freshwater that has in Africa countries the 43 = eastern S = southern Africa, Africa.” 182 CHAPTER 6 | FRESHWATER CRABS Africa, E = eastern Africa, N = northern Africa. Africa,N=northern Africa, E=eastern occurrence, andasummaryoftheirconservationstatus. Table 6.2.Thethreatened speciesoffreshwater crabsfoundincontinentalAfrica,withtheirregions of andfreshwaterpatterns ecoregion boundariesfoundinthe no closecorrelation betweenfreshwater crabdistribution et al Temperate freshwater ecoregions) (Thieme andCoastal,Cuanza,Zambezi,Southern Eastern Upper Guinea,West CoastEquatorial,Congo,Great Lakes, freshwater ecoregions (e.g.,theNorthAfrican,Nilo-Sudan, crabs are alsorepresented inmostofthecontinent’s representation inAfrica’s aquaticecosystems,freshwater and drainforested highlands.Becauseoftheirwidespread Africa, especiallythosethatfl highest intheriversandstreams ofthemajorriverbasins freshwater resources aboveground. Speciesdiversityis savanna rivers,andevenxericsystemswhere there are mountain systems,large lakes,large riverrapids,dry moist forest rivers,Mediterraneansystems,highlandand continental Africa,includingfl Freshwater crabsare foundinallmajorhabitattypes 6.1.1 Biogeographicpatterns 2 1 7 6 5 4 3 8 9 10 11 14 13 12 18 17 16 15 19 20 21 22 23 24 26 25 . 2008).However, withonlyafewexceptions,there is Species Liberonautes lugbe Liberonautes grandbassa Louisea balssi Liberonautes rubigimanus Liberonautes nanoides Globonautes macropus Afrithelphusa monodosa Louisea edeaensis Potamonautes gonocristatus Potamonautes idjiwiensis Potamonautes mutandensis Potamonautes choloensis Liberonautes nimba Potamonautes platycentron Potamonautes lividus Potamonautes infravallatus Potamonautes ignestii Potamonautes gerdalensis Potamonautes montivagus Potamonautes pilosus Potamonautes raybouldi Potamonautes reidi Potamonautes triangulus Potamonautes unisulcatus Potamonemus sachsi Potamonautes xiphoidus ow through lowlandrainforest oodplains, swamps,lakes, et al . 2005;Abell itiuinRdLs aeoyRedListCriteria RedListCategory Distribution Regional / UB1ab(iii)+2ab(iii) VU W/C / NB1ab(i,iii)+2ab(i,iii) EN C/E / NB1ab(iii,v)+2ab(iii,v);C1+2a(ii) EN C/E RB1ab(iii)+2ab(iii) CRB1ab(iii) CR W W NB2ab(iii) B1ab(iii)+2ab(iii) B1ab(iii)+2ab(iii);C2a(i) B1ab(iii)+2ab(iii);C2a(i) EN EN EN EN W W W W UB1ab(iii)+2ab(iii);D2 VU W UB1ab(iii)+2ab(iii) VU W UB1ab(iii)+2ab(iii) VU W NB1ab(i,ii,iii,iv,v) EN C NB1ab(i,ii,iii,iv,v) EN C NB1ab(i,ii,iii,v) EN C UB1ab(iii)+2ab(iii) VU S UB1ab(i) B1ab(iii)+2ab(iii) VU EN E E UB1ab(i);D2 B1ab(i);D2 B1ab(i);D2 VU VU VU E E E UB2ab(i) VU E UB1ab(i,iii);D2 VU E UB2ab(iii) VU E UB1ab(i,ii,iii);D2 VU E UB1ab(iii)+2ab(iii);D2 VU E C = central Africa, W = western Africa, S = southern Africa,S=southern C=centralAfrica,Wwestern species thathavearestricted distribution. distribution coincideswithecoregion boundariesare those African region. Theonlyinstanceswhere freshwater crabs least twofreshwatercrabsspecies. The WhiteVolta Riverasanexampleofsuitablehabitatforat

FRESHWATER CRABS

The Purple March Crab Afrithelphusa monodosa (Endangered) which lives in swamps and year-round wetland habitats in north- western Guinea where it is known from only a few specimens from two localities. This species is clearly a competent air-breather and has a pair of well-developed pseudolungs. It is mainly threatened by habitat loss and degradation. © PIOTR NASKREKI

6.2 Conservation status with 26 (21%) of the 120 species assessed as globally Threatened (Table 6.2 and 6.3, and Figure 6.1a). There is The conservation status of Africa’s freshwater crab fauna insuffi cient information to assess the status of 27 species was assessed by Cumberlidge et al. (2009) using the which were categorised as Data Defi cient (DD) due to a lack IUCN Red List Categories and Criteria at the global scale specimens, and locality and population data (Cumberlidge (IUCN 2003), and individual species assessments are et al. 2009; Table 6.3, and Figure 6.1a). If all Data Defi cient freely available through the IUCN Red List website (www. (DD) species also proved to be Threatened, the level of iucnredlist.org). Results reveal current high levels of threat threat could be as high as 44%.

Figure 6.1. The proportion (%) of freshwater crab species in each regional IUCN Red List Category in mainland continental Africa (Cumberlidge et al. 2009, Appendix 1): a) all 120 species from the region; and b) following removal of the 27 DD species.

183 184 CHAPTER 6 | FRESHWATER CRABS species offreshwater crabsfrom Africacouldbeconfi species (3%)were assessedasNearThreatened (NT).No (VU) (14species)(Figure 6.1bandTable 6.3).Justthree (two species),Endangered (EN)(10species)orVulnerable threatened categories,eitherasCriticallyEndangered (CR) of the94assessedspecieswere listedinoneofthree species of eight speciesof eight oftheninespecies Potamonemus of Cumberlidge streams intheforested partsofAfrica(Cumberlidge1999; of theseliveinrivers,marshylowlands,ormountain and were (LC),andmost judgedtobeLeastConcern (65 species,69%,Figure 6.1b)belongtosevengenera Of the94speciesthatcouldbeassessed,majority found in both C + S). C = central Africa, W = western Africa, S = southern Africa, E = eastern Africa, N = northern Africa. Africa,N=northern Africa,E=eastern Africa,S=southern found inbothC+S).=centralAfrica,Wwestern both E+C,E+S,andSCE);ENincludestwospeciesthatare foundinbothC+E);DDincludesonespeciesthatis fi with an*=totalspeciesperregion includingthosespecies thatare foundinmore thanoneregion. Forexample,LCincludes Appendix 1). Table 6.3. Aubry’s Crab of breathing air. during rainstorms itisoftenfoundonlandbecause it iscapable and ponds,digs burrowsnearwaterways;atnight and Equatorial Guinea.Thislargespecies inhabitsstreams,rivers, border oftheCampoMaanNational Parkontheborderto Central Africa.Thisspecimenwas collected onthewestern widespread speciesfoundthroughout muchofwesternand ve speciesthatare foundinbothW+C,twospeciesthatare foundinbothE+N,andonespecieseachthatis UNRdLs aeoyNWCES E C W N IUCN RedListCategory es ocr 01 41 64 14 2 10 2 15 1 0 0 24 9 2 16 4 1 0 0 2 10 3 0 0 4 2 0 0 0 0 2 0 Least Concern Near Threatened Vulnerable Endangered Critically Endangered aaDficet0 25 25 117 4 5 12 20* 44* 5 31* 0 24* 2* Total speciesperregion* Data Defi cient Potamonautes

The numberofAfricancrabspeciesineachIUCNRedListcategorybyregion (Cumberlidge Sudanonautesaubryi Erimetopus The totalnumberofspeciesperCategorydoesnotincludefoundinmore thanoneregion. Numbers et al , eightofthe11species © JENSKIPPING are LC,asare oneofthethree speciesof Liberonautes . 2009). and

Forty-three outofthe76species Potamon Platythelphusa , andoneeachofthetwo (Least Concern)isa . Twenty-six (28%) Sudanonautes , three ofthe rmed REGION , (Smith Africa Africa (CumberlidgeandDaniels2008),western Africa(Darwall available foreastern Detailed assessmentsofspecies’regional statusare also Extinct untilexhaustivesurveyshavebeencarriedout. be notedthataspeciescannotformallyassessedas Extinct (EX)orintheWild(EW).However, itshould central Africa(Brooks species numbers (Cumberlidge 1999). For example,there identical forest areas that haveagradientofdeclining areas of highbioticdiversitysurrounded byapparently shown bythedistributionofspecies inAfricanforests is the fewestinsavannaecosystems. Oneinteresting pattern rainforest ecosystems, especiallyinhighlandareas, and of water, withthehighest numberofspeciesoccurringin appears todependonvegetation coverandtheavailability in highlandregions (Cumberlidge1999).Speciesdiversity are noticeablymore abundantintherainforest, especially found inallofthemajorecosystemscontinentalAfrica,but and vegetationcover. Forexample,freshwater crabsare region, andvarieswithecosystem,aquaticdrainagebasins, crab faunainAfricaisnotuniform,changesfrom region to available dataindicatethatthecompositionoffreshwater are necessarytoascertaintheiractualdistributions.The just afewrecords, andinthesecasesfurther collections that are knownonlyfrom eitherthetype locality orfrom African speciesare quitewellstudied,there are stillsome are stilllikelytobeincomplete.Althoughamajorityofthe have beenderivedfrom allavailablespecimenrecords but (Cumberlidge 1999;Table 6.1).Distributiondatausedhere endemic tothecontinent,anddistributedover43countries recognisable freshwater crabfauna,withallspecies Continental Africahasarich,highlydiverse,anddistinctly ofspeciesrichness 6.3 Patterns et al . 2009), northern Africa(Garcia. 2009),northern Red ListCategory et al Total speciesper . 2011). et al . 2005), southern . 2005),southern et al et al . 2010),and . 2009, . 2009, CHAPTER 6 | FRESHWATER CRABS 185 are are . 2000), is found et al Erimetopus has a number of Deckenia REGION 6641 and Potamon Louisea DENIS TWEDDLE © 9 00990 8 07000 4 04000 3 01200 2 00020 2 00200 2 00200 1 10000 1 01000 N = northern W = western Africa, 76 2 4 23 33 19 1105510 Potamonautes Genus # Species N W C E S Sudanonautes Platythelphusa Liberonautes* Afrithelphusa* Potamonemus Deckenia* Erimetopus* Louisea* Potamon Globonautes* Genera /region 2 endemic to western Africa, both found only in central Africa, and only in eastern Africa. Although within Africa this genus is found only in the Palaearctic the Palaearctic within Africa this genus is found only in zone of northern Africa. Table 6.4. Number of species and genera found in found and genera of species Number 6.4. Table in Africa. region each central Africa, E = easternAfrica, C = S = southern Africa, endemic. regional Africa. * = and Middle wide distributional ranges in the Mediterranean East as far east as the Himalayas (Brandis

, and are all are For example,

Sudanonautes Potamonautes Platythelphusa Liberonautes include some widespread include some widespread ve regions, and (Least Concern) is a relatively abundant and widespread species found in large slow fl owing rivers in (Least Concern) is a relatively abundant and widespread species found in large slow fl Liberonautes occur in two regions. occur in two regions. , and is found in all fi ve African regions has genera with species that ve African regions Potamonautes lirrangensis rainforests across central and eastern Africa. It is subject to a small commercial fi shery in Malawi. rainforests across central and eastern Africa. It is subject to a small commercial fi Afrithelphusa, Globonautes are three relatively small areas of the African rainforest that rainforest African of the small areas relatively three are of species number high an unusually out as having stand the of endemic species: than normal number and a higher in western Africa, the Lower Guinea forest Upper Guinea of eastern and the forests Nigeria and Cameroon, in forest (Cumberlidge 1999). D. R. Congo taxonomic diversity is highest in At the genus level the western in eastern and central Africa (six genera), lower and lowest in northernAfrica (four genera), Africa (two 6.4). genera) and southern (one genus) (Table Africa (with Although eastern region Africa is the most species rich western genera and two subfamilies), 44 species in three the most diverse (with 24 species, Africa stands out as 6.4). Distribution subfamilies) (Table six genera and two at the genus level indicate that each patterns considered of the fi that have have a wide distribution, and many other species Five out of eleven genera have a distribution. a restricted For example, than one region. wide distribution in more Potamonautes species, and some species that are endemic to a particular species, and some species that are restricted In contrast, six genera have a relatively region. endemic to a particular region. range and are Sudanonautes is found in three regions, while regions, is found in three Potamonemus 186 CHAPTER 6 | FRESHWATER CRABS Stewart 1997;Corace et al species inAfrica(Bott,1955,1959,1969,1970;Stewart Potamonautes 6.3.1 western Africa; the western limitofthegenus isSenegal, Africa;the western western The eightspecies of 1999; CumberlidgeandBoyko2000). have arestricted distribution incentralAfrica(Cumberlidge the othertwospecies, are widespread throughoutandcentralAfrica,while western Sudan savanna.Eightofthe11 speciesof monodi savanna inNigeria,whilethesemi-terrestrial air-breathing savanna. and occur exclusivelyinrainforest habitats,while granulatus, S.orthostylis,chavanesii rivers) andonland. most aquatichabitats(standingwater, streams andmajor rainforest, GuineaandSudansavanna),are foundin andcentralAfrica(tropical major ecosystemsofwestern Bioko. Speciesof Nigeria, south-westCameroon, andontheislandof species 1999 (Cumberlidge 1989,1991,1993b,c,1994b,1995,a,b,c,d, Africa as fareastUgandaandSudanineastern and centralAfrica,withonespecies( Sudanonautes Niger Riverfl is evenfoundintheSaharadesertMali,butonlywhere the along theNileRiver).Onespecies( Africa(bothwidespread and twoinnorthern speciesfound species of (33, 24,and20speciesrespectively), andthere are onlyfour Potamonautes forested CongoRiverbasin.Thevastmajorityofspecies species of are endemictoaveryrestricted area. Thelargest numbersof breathing habit,whileothershaveanarrow distributionand the continent,somehaveadoptedasemi-terrestrial air- and are associatedwiththemajorriverbasinsthroughout region. Somespeciesof (the Maghreb) thatliesinthePalaearctic zoogeographical does notoccurisinthecoastalfringeofnorth-westAfrica Africa (Figure 6.2).Theonlyregion where from theCapetoCairo, andfromof SenegaltotheHorn Africa, andare distributedthroughout sub-SaharanAfrica species ofthislarge genusare foundinallfi and Tavares 2006;CumberlidgeandDobson2008).The76 2004, 2006;CumberlidgeandBoyko2000; Cumberlidge andVanini 2004;ReedandCumberlidge S.fl ) ( . 1995; Cumberlidge 1997, 1998, 1999, 2009a,b; . 1995;Cumberlidge1997,1998,1999,2009a,b; distribution patterns All freshwater crabspecies:interpretation of Figure 6.2 istheonlyspeciesfoundinbothGuineaanddry

oweri are foundintheforested region ofsouth-east Sudanonautes kagoroensis Potamonautes Potamonautes ows through Timbuktu (Cumberlidge 1999). are foundinbothrainforest andwoodland includes 11 species distributed in western includes11speciesdistributedinwestern includesbyfarthelargest numberof occur in eastern, central, and southern Africa Africa central,andsouthern occur ineastern, ) . Thelargest numbersof Sudanonautes africanus, S. nigeria, S. Sudanonautesafricanus,S.nigeria, Sudanonautes Liberonautes et al S. orthostylis occur in eastern Africa and in the Africaandinthe occurineastern in western Africa (all endemic), Africa(allendemic), inwestern . 2001;Cumberlidge Potamonautes occur in most of the occurinmostofthe Potamonautes ecorssei and occurs only in Guinea occursonlyinGuinea are all restricted to S. ﬂ and S. sangha oweri are widespread Potamonautes Sudanonautes Sudanonautes Sudanonautes ve regions of S. faradjensis et al ) reaching S. aubryi S. aubryi . 2002; . 2002; , each S. S. )

© population growthandindustrialagrariandevelopment. threats suchashabitatlossanddegradationlinkedto Tunisia. ItisLeastConcernandaffectedbyhumaninduced endemic tothestreamsandriversofMorocco,Algeria The freshwatercrab Africa. partoftheUpperGuineaforestin thewestern of 1996a,b; 1999).Thefourspeciesof forest (CumberlidgeandSachs1991;Cumberlidge1991, in smallwaterreservoirs intree holesin closed canopy of theUpperGuineaforest inLiberiaandGuinea,lives the LiberianTree part HoleCrab,occursonlyinthewestern 1992; Cumberlidge1993a;1999). leave thewateratnighttofeed(CumberlidgeandClark 6.2). Thesespeciesoccurinsmallstreams andprobably Nigeria, south-westCameroon,Togo andwestern (Figure and centralAfricaintheforested highlandsofsouth-east The three speciesof nanoides nimba L. paludicolis are foundinaquatichabitats(streams ormajorrivers)while in bothGuineaandSudansavannazones.Mostspecies mostly inrainforest, although and Côted’Ivoire. Speciesof species eachhavealimiteddistributioninLiberia,Guinea, entire rangeofthegenus,whileremaining seven 6.2). Onespecies( (Cumberlidge 1999;CumberlidgeandHuguet2003 limitisMali limitisGhana,andthenorthern the eastern found onlyinGuinea,while 1996a,b; 1999).Thetwospecies of only knownfrom SierraLeone(Cumberlidge1987,1991; other species, is relatively widespread inthelowerCongoRiver, whilethe 1999). Oneofthetwospecies of and themoistlowlandrainforests (Cumberlidge1994a, only insouth-westCameroon intheforested highlands al distribution (Cumberlidge andReed2004;Cumberlidge . 2009). All ninespecies of JEAN-PIERRE BOUDOT are foundathighaltitudes,and Afrithelphusa gerhildae are foundexclusivelyinthemajorrivers. ismore terrestrial and E. vandenbrandeni, Potamonalgeriense L. latidactylus Potamonemus Platythelphusa A. afzelii L. latidactylus and ) iswidespread overthe Liberonautes Globonautesmacropus Erimetopus Afrithelphusa occur in only in western occurinonlywestern is more restricted inits A. monodosa L. rubigimanus and (Least Concern)is Louisea L. chaperi A. leonensis are endemicto isalsofound ( are found are found E. brazzae) occur only are both ) ( and and Figure are et L. L. ,

CHAPTER 6 | FRESHWATER CRABS 187

) ), ). ), four P. lividus P. L. balssi P. idjwiensis, P. and P. choloensis, P. choloensis, P. P. P. xiphoidus P. P. gonocristatus P. and and L. edeaensis P. reidi, P. triangulus, Potamonemus reidi, P. P. ), one from southern), one from Africa ( ), four from eastern), four from Africa ( A. monodosa, G. macropus, L. nanoides, L. nimba and L. rubigimanus central Africa (the most speciose region) and just one central Africa (the most speciose region) no threatened are in southern 6.3). There Africa (Figure species in the northern Of the threatened Africa region. from assessed as EN, of which four are species, 10 were western Africa ( and platycentron, mutandensis, P. P. Africa ( central and two from southern None of the species from Africa was assessed as assessed as VU, EN. A further fourteen species were eastern from Africa ( of which nine are montivagus, P. infravallatus, P. ignestii, P. gerdalensis, P. unisulcatus, raybouldi, P. pilosus, P. from western Africa ( from sachsi, et

2005). One . 2009; Figure . 2009; Figure are endemic to are et al. et al . 1999; Marijnissen ) is found in continental Deckenia et al . 1995; Marijnissen P. algeriensis P. ( et al . 2000; Cumberlidge 2009a). Potamon et al 2004), while both species of Lake Tanganyika (Cumberlidge Lake Tanganyika al. eastern Africa (Ng species of 6.3.2 Threatened species 6.3.2 Threatened Some 28% (26 out of 94 species) of African freshwater threatened as being in one of the three assessed crabs were categories (CR, EN, VU) (Cumberlidge Africa and is endemic to the Mahgreb in northern Africa and is endemic to the Mahgreb Africa (Brandis found in 6.5). Most of these (20 species) are 6.1a; Table either eastern Africa (11 species) or western (nine Africa species in threatened only three are species), and there Figure 6.2. The distribution of freshwater crab species across mainland continental Africa. Species richness = richness Africa. Species continental mainland across species crab of freshwater distribution 6.2. The Figure sub-catchment. per river/lake of species number 188 CHAPTER 6 | FRESHWATER CRABS IUCN RedListCategory. Table 6.5.Numberofspeciesfrom eachgenusin richness =numberofspeciesperriver/lakesub-catchment. Figure 6.3. eu CN UE RDD CR EN VU NT LC Genus oa pce 431 0227 2 10 14 3 64 Total species Globonautes Potamon Louisea Erimetopus Deckenia Potamonemus Afrithelphusa Liberonautes Platythelphusa Sudanonautes Potamonautes

The distributionofThreatened freshwater crabspeciesacross mainlandcontinentalAfrica.Species 430124 018 000100 010000 000200 100001 020000 101001 000103 301220 800001 800003 IUCN REDLISTCATEGORY

a lackofknowledgeorunder-collection. Thesespeciesare are notsimplyaproduct ofomissionerrors stemmingfrom Africa (Figure 6.4).Thelimiteddistributionsofthesespecies restricted rangespeciesare irregularly distributedthroughout western Africa).Twowestern species( and onefrom centralAfrica( have arestricted range(<20,000km Excluding DDspecies,some29specieswere found to 6.3.3 Restrictedrangespecies habitat degradationandpollution. and agriculturaldevelopmenttheassociatedaquatic threats tothesespecieswere identiﬁ (both fromAfrica)were eastern assessedasNT. Themain Africa. Justtwospecies, were assessedasCR,bothofwhichare from western

Deckenia imitatrix P. sachsi L. grandbassa 2 ed asurban,industrial, ) (Table 6.6),andthese , which is also found in , whichisalsofoundin and and L. lugbe D. mitis )

CHAPTER 6 | FRESHWATER CRABS 189 WINKS EMMERSON © (Vulnerable) is a (Vulnerable) Potamonautes lividus relatively restricted range species endemic to swamp forests in north-eastern Kwa-Zulu Natal, South Africa. The Blue River Crab (and 2 or less). 2 LC < 20,000 <10 N C LC < 800 <5 N C LC < 20,000 8 N C LC < 5,000 7 N W VU < 20,000 <10 N C VUVUVUVUVU < 20,000VU < 20,000VU < 20,000VU < 20,000VU 4 < 20,000VU <10 < 20,000VU <5 < 20,000VU <5 < 20,000 YVU <10 N < 2,000 <10 < 20,000 N <10 < 20,000 N <5 N < 20,000 W E <10 Y < 2,000 <10 E N E 6 N E <5 N S <5 Y E N E N E N W W E E ENENENENEN < 5,000 < 5,000 < 5,000 < 20,000 2 < 5,000 5 1 3 N 2 N N Y W N W W W C EN < 5,000 3 N C ENENENEN < 5,000 500 < 500 < 500 < <10 <5 <5 <5 N N N N E E E E CRCR ~ 100 100 ~ 1 1 N N W W RL = IUCN Red List status; LC = Least Concern; VU = Vulnerable; EN = Endangered; CR = Critically = Critically CR = Endangered; EN Concern; LC = Least List status; IUCN Red RL = VU = Vulnerable; cient. cally recorded, through many surveys conducted through cally recorded, Liberonautes lugbe Afrithelphusa monodosa Globonautes macropus Liberonautes nanoides Liberonautes rubigimanus Louisea edeaensis Liberonautes grandbassa Species RL Category Range (km2) # Loc PA Region Louisea balssi Potamonautes gonocristatus Potamonautes idjiwiensis Potamonautes mutandensis Potamonautes platycentron Erimetopus brazzae Potamonautes margaritarius Potamonemus mambilorum Potamonemus sachsi Sudanonautes kagoroensis Liberonautes nimba Potamonautes choloensis Potamonautes gerdalensis Potamonautes ignestii Potamonautes infravallatus Potamonautes lividus Potamonautes montivagus Potamonautes pilosus Potamonautes raybouldi Potamonautes reidi Potamonautes triangulus Potamonautes unisulcatus Potamonautes xiphoidus Despite the dangers of population fragmentation, current Despite the dangers of population fragmentation, current population levels of stenotopic species assessed as LC or to be stable because they have been estimated NT were Table 6.6. Species of freshwater crabs of Africa restricted to single river basins, excluding those considered to considered those excluding river basins, to single Africa restricted crabs of of freshwater 6.6. Species Table Defi be Data Number of specimens; #Loc = based on all known distribution range of the species Range = an estimation Endangered; of N = no; regions Y = yes, area; = found in a protected collected; PA the species was which localities from discontinuous noted as N = northern Africa, W = westernAfrica are E = eastern C = central Africa, Africa, = southern Africa, S Africa. specifi they may over the years, as being absent in localities where Any disruption to the habitats have been expected to occur. development, pollution, or of these species (either from have serious consequences, given that could political unrest) range species have been assessed as 24 of these restricted range is 6.6). Any species with a restricted (Table Threatened population fragmentation potentially vulnerable to extreme rapid decline, and even extinction, in a a and could suffer should dramatic changes in land-use short time relatively of immediate concern habitat. It is therefore its suddenly affect that 24 of the 94 crab species that could be assessed are ranges of less than 20,000km distribution known from some of these have an estimated range of 5,000km 190 CHAPTER 6 | FRESHWATER CRABS are from centralAfrica ( be DD(Table 6.3, Figure 6.1a).Thirteen oftheDDspecies Some 27ofthe 94 species(28%)assessedwere judgedto 6.3.4 DataDefi al species nowindecline(Cumberlidge 2008;Cumberlidge their beingtheremnant populationsofformerlywidespread (marginal) habitatorthrough islandcolonization,ratherthan relatively recently inresponse toisolationinaspecialised unknown, butitisthoughtlikelythattheyhavespeciated the restricted rangesofthestenotopicspeciesare largely and endangertheirlong-termexistence.Thereasons for threats thatwouldimpactthehealthofthosestreams collected recently andthere are noidentifi mainland continentalAfrica.Speciesrichness=numberofspeciesperriver/lakesub-catchment. Figure 6.4.Thedistributionoffreshwater crabspecieswithseverely restricted distributionalrangesacross . 2009). cient species E. vandenbrandeni, P. acristatus, able immediate immediate able et et and schubotzi, P. semilunaris, Potamonemusasylos,S.orthostylis P. adeleae,P. adentatus,P. lueboensis,P. punctatus,P. in viewof theabsenceof recent informationon theirExtent found inrecent years.Thesespecies have beenlistedasDD thatinmanycases theseDDspecieshavenotbeen concern Cumberlidge 1999; the distributionoftheselittle-known species(Bott1955; available, ascarcity that continuestofueluncertaintyabout of DDspeciesrefl Africa were foundto be DD.Therelatively highproportion macrobrachii three are fromAfrica( southern bipartitus, P. didieri,P. rodolphianus, are fromAfrica( eastern gerhildae, A.leonensis,P. senegalensis, S.sangha ) (Figure 6.5).Noneofthespeciesfrom northern ), fi ve are fromAfrica( western ects thegenerallackofspecimens P. praelongata,P. P. amalerensis, P. kensleyi,P. dubius, et al and . 2009).Itisofgreat P. rothschildsi and

S.nigeria), A. afzelii, A. A. afzelii, and ), and ), and P. six six

CHAPTER 6 | FRESHWATER CRABS 191 owing 6.4 Major threats 6.4 Major threats crabs include water African freshwater to The main threats pollution, urban, industrial, and agricultural development, human from resulting and habitat loss and deforestation population and urban and agricultural expansion (Figure 6.6). 6.4.1 Habitat destruction species include habitat destruction to the endemic Threats driven by timber extraction, in the form of deforestation the demands of increasing agriculture, mining, increasing industrial development, the alteration of fast fl and the power, of hydroelectric rivers for the creation drainage of wetlands for farming and other uses (Figure E. S. cient and cient freshwater crab species across mainland continental Africa. Species Species Africa. continental mainland across crab species cient freshwater P. schubotzi, S. orthostylis P. or where they are known from either only known from they are or where , ) ) and the full range extent is uncertain. It is possible . 2009). It is also of concern that many of these species of Occurrence, habitat, ecological requirements, population population ecological requirements, habitat, of Occurrence, (Cumberlidge and long-term threats size, population trends, et al a few individuals collected many years known only from are ago, and that no new specimens have been found recently. is insuffi there The DD status is also assigned where information either on their taxonomic distinction (e.g. vandenbrandeni sangha that in some cases the DD status may be due to under- sampling but, as mentioned above, this is not thought to is be the case for many of the DD species. Further research needed on all of these species because, at least, they may range endemics vulnerable to habitat to be restricted prove loss. one or only a few localities ( Figure 6.5. The distribution of Data Defi of Data distribution 6.5. The Figure sub-catchment. per river/lake species = number richness 192 CHAPTER 6 | FRESHWATER CRABS Figure 6.6. towns inAfricaincludepollutionbysewageandindustrial aquatic communitiesinriversassociatedwithcitiesand to freshwater ecosystems.Potentialfuture threats to and soilerosion thatcontributesenvironmental damage movements, whichare oftenaccompaniedbydeforestation habitat destructionincludepoliticalunrest andrefugee the forest. Otherthreats thatresult indeforestation and access toremote andpreviously undisturbedpartsof is furtherexacerbatedbyloggingroads thatprovide 6.6). Destructionoftheforests inmanypartsofAfrica region of theUpperGuinean Rainforest. Deforestation near thevillageSimpainwesternGhana, inthe

Major threats tofreshwater crabspeciesinmainlandcontinentalAfrica. © JOHANNES FÖRSTER egrets, herons, giant kingﬁ necked otters, water mongooses,Africancivets, kites, A large numberofAfrican predators –including yellow- 6.4.3 health ofpeoplethatmayeatthem. metals incrabscouldposeanincreasing problem forthe in theseareas, notleastbecausethebioaccumulationof should begiventotheimprovement ofthewaterquality contain highlevelsofcontaminants.Immediateattention feeders thatingestotherinvertebratesanddetritusmay freshwater crabpopulationsbecausetheyare benthic rivers andotherfreshwater bodies.Thesepollutantsimpact sewage systemsinurbanareas inAfricacanaccumulate iron ore, andcoltan,from organic wastesfrom leaking Pollutants from miningactivitiesfordiamonds,gold,bauxite, 6.4.2 Pollution survival. the future willhavethebiggestimpactontheirlong-term management ofAfrica’s forests andwaterresources in to range-restricted endemicspecies,andthecareful factors combinetoincrease theoveralllevelofthreat more research needstobecarried out.Alloftheabove farmers mayprove tobelethalfreshwater crabs,but and generalwaste.Someagriculturalpesticidesusedby introduced species Natural predators and competitionwith shers, monitorlizards, CHAPTER 6 | FRESHWATER CRABS 193 et S. cation and c ecologies and P. lirrangensis P.

© RANDALL BRUMMETT ed and taxonomic advances ed and taxonomic P. perlatus, P. that are all currently assessed as LC primarily all currently that are cant areas of this vast continent still remain of this vast continent still remain cant areas ciently explored, and new species of freshwater and new species of freshwater ciently explored, . 2002). keys. Although taxonomic knowledge has advanced and museum collections of considerably recently, work deal of a great have improved, crabs freshwater is a need for further surveys still needs to be done. There Signifi insuffi in if collection efforts to be discovered sure crabs are intensifi are areas remote available in the form of identifi readily become more recently, some taxa currently assumed to be widespread assumed to be widespread some taxa currently recently, to be complexes of several and common may prove distinct cryptic taxa, each with specifi conservation action. Three direct distributions requiring such possibilities are actions required 6.5 Research granulatus on account of their wide distributional ranges. However, the distribution patterns of these species consist of many deal subpopulations that show a great isolated relatively of morphological variation, and further investigations may show these species to be species complexes (Daniels al

is rapidly Procambarus Platythelphusa sh ( sh (Foster and Harper Procambarus clarkii . 2009). The ecological importance of et al ) in the rivers of western Kenya, which out-competed clarkii crab in freshwater native crabs. The drop the and replaced populations in these systems led to a subsequent decline in clawless otter populations that fed on crabs, most competition with other predators likely due to increased attracted by the abundance of crayfi indicate that 2006, 2007; Ogada 2006). Recent reports the highly competitive in Eastern Africa and may even reach elsewhere spreading of populations it could threaten where Lake Tanganyika, in the endemic genus crabs freshwater pers. comm.). (Howard, issues 6.4.4 Taxonomic crabs may prove The evolving taxonomy of freshwater to be a challenge for conservation planning in the future has been made because, although a lot of progress and crocodiles – depend on freshwater crabs as vital – depend on freshwater and crocodiles crabs freshwater components of their diet. This is because in African aquatic macro-invertebrates the largest are the food chain ecosystems and form an integral part of (Cumberlidge in food webs in Africa is underlined by crabs freshwater the impact of alien North American crayfi Pollution from urban and domestic sources is a major threat to threatened freshwater crab species. Pollution from urban and domestic sources is The Louisiana Crayfish Procambarus clarkia has become seriously invasive throughout inland waters in Africa. It is omnivorous and will hunt and compete with the native freshwater crab species. © GEOFFREY HOWARD CHAPTER 6 | FRESHWATER CRABS 195 ne line rst step . 2009) represents a fi . 2009) represents et al c habitat requirements and a restricted and a restricted requirements c habitat cation of threatened species within the cation of threatened eld survey. The scarcity of available specimens of available scarcity The eld survey. cient fi cient toward the identifi toward the development of a conservation and toward region range of strategy for endemic species. The restricted many species, together with the on-going human-induced primary are loss of habitat in many parts of the region, causes of concern for the long-term survival of this fauna. of endemism, crabs have a high degree freshwater Africa’s with many species living in specialised habitats such as highlands, and river rapids, lowland marshes, forested to determine is recommended islands. Additional research areas size and design of protected the minimum effective such as crabs. species for freshwater and streams. The increasing loss of natural vegetation and vegetation of natural loss The increasing and streams. agriculture and land development of as a result pollution rivers in the the lowland affect likely to however, are, that aquatic species and many of the wholly long term, species be vulnerable. Even could eventually live there should declines catastrophic LC could suffer assessed as land development, hydrology, abrupt changes in be there how the highland It is not known or pesticide-use regimes. disturbance and pollution but, taxa will cope with habitat is it habitat requirements, considering their specialised as species will not adapt as readily likely that most of these lowland species. In many countries widespread the more development, often only a fi with a rapid pace of one assessed assessed as LC from separates a species is assessed as EN. one that from as VU, or a VU species impact on could have a dramatic Development projects species with specifi be therefore distribution. Conservation activities should the integrity of sites and aimed primarily at preserving monitoring key habitats while at the same time closely crab populations. freshwater crabs judged to be The 27 species of African freshwater of assigned to this category primarily as the result DD were insuffi the long-term is in some parts of the continent due to little is known of poor security situation, and as a result to these or threats the habitat needs, population trends, it is information has been gathered, species. When more have a relatively expected that almost all DD species will distribution and be endemic to the river basin restricted found. they are where crabs in The conservation assessment of freshwater Africa (Cumberlidge et ed to c threats to Africa’s to Africa’s c threats . 2008; Cumberlidge consider shery managers ne species distributions, distributions, ne species et al cally include the conservation and cally include the . 2009). et al c habitat requirements, describe population population describe requirements, c habitat ne specifi . 2009). 6.6. Conservation recommendations 6.6. Conservation patterns crabs The biology and distribution of the freshwater the potential as are becoming better known, of Africa are 27 species (28%) to their long-term survival. With threats Africa crabs from of freshwater of the 94 non-DD species global extinction, assessed as being at risk of currently endemic largely of the continent’s the long-term survival crab fauna is a concern. it is freshwater Nevertheless, plans for threatened hoped that conservation recovery identifi species will be developed for those species this Red List be in need of conservation action through (Collen assessment process al crabs The conservation of many species of freshwater of natural of areas depends primarily on preservation Although it maintain water quality. enough to habitat large freshwater is not yet known exactly how sensitive African is evidence to polluted or silted waters, there crabs are not likely to survive Asia that similar crabs are from 2007). and Yeo when exposed to these factors (Ng and exploitation of natural Development, agriculture in developing economies, necessary realities are products crab may have to be made if freshwater but compromises Judicious not to be extirpated in the future. species are is unlikely to cause species use of resources and careful not heavily are extinctions as long as water drainages and vegetation some forest polluted or redirected, respected are areas cover is maintained, and protected (Cumberlidge sustainable use of local populations of river crabs. use of local populations sustainable Common species assessed as LC have a wide distribution of the region in the rivers, wetlands, and mountain streams tolerant of changes to be relatively have proved and, so far, aquatic ecosystems. It is encouraging in land-use affecting species can persist in the already adaptable that these more disturbed and visibly polluted parts of the lowland rivers to discover new species, refi species, new to discover specifi and identify trends, levels and to the crab fauna. It is vital unique freshwater important and fi ecosystems that health of these that specifi measures defi 196 CHAPTER 6 | SPECIES IN THE SPOTLIGHT C UNDP/GEF Project on LakeTanganyika, Regional CoordinationUnit, c/oSecretariat oftheLakeTanganyika Authority, Route Pri ¹ \mfclk`feXe[\okèZk`fe èCXb\KXe^Xep`bX1Xdf[\c]fijkl[`\jè 8le`hl\jg\Z`\jÕ Platythelphusa maculata crevices underneathrocksandstones. that enablesittohideinsidenarrow has evolvedanextremelyfl shells. claw thattheyusetocrushsnail with onesignifi of instance, bothmalesandfemales al. adaptive evolution(Cumberlidge them idealsubjectsforstudiesin and ecologicalfeaturesmaking a strikingvarietyofmorphological Africa. Platythelphusidcrabshave species fl form theonlyknownlacustrine the genus platynotus crabs, including derived from freshwater ancestors. derived fromfreshwater ancestors. that allLakeTanganyika crabsare However, geneticevidenceindicates millions ofyears(Moore1903). sea andremainedunchangedover in thelakeafterarecessionof of marinecrabsthatgottrapped platythelphusids werearelictgroup Initially, itwasthoughtthatthe Glor 2010,andreferencestherein). processes ofadaptiveevolution(e.g. famed naturallaboratoryforstudying diverse lake,LakeTanganyika isa million years)andmostspecies tanganyicensis. large endemicsnail refuge withinemptyshellsofthe species, whichisoftenfoundtaking other hand,isatinyround-bodied Platythelphusaarmata As Africa’s oldest (estimated 9-12 As Africa’soldest(estimated9-12 1999;Marijnissen Bujumura, Burundi Platythelphusa conculcata endemic species of freshwater endemic speciesoffreshwater Lakes inharbouring10 among theAfricanGreat ake Tanganyika isunique Jg\Z`\jèk_\jgfkc`^_k ock offreshwatercrabsin Platythelphusa andninespeciesin cantly enlarged Potamonautes Neothauma et al. ,

on the on the which which are armed are armed 2004).For at body et 2000). Research on the crabs shows 2000). Research on thecrabsshows niche diversifi result ofcompetitionandecological in theAfricanGreatLakesare the highlevelsofspeciesdiversity longstanding hypothesisisthat relatively shortperiodoftime.A seem tohaveevolvedwithina between theplatythelphusidcrabs million yearsago. during thesameperiod,about3.3-2.5 (Day andWilkinson 2006),diverged the et al. (Duftner including severalcichlidgroups Marijnissen basins (CaneandMolnar2001; Tanganyika wassplitintoseparate dropped substantiallyandLake and drought,whenthelakelevels with aperiodofclimatechange the platythelpusidscoincides The evolutionarydivergenceof Potamonautes platynotus out ofthewatertofeedamongstrocksalonglakeshore. The morphological differences The morphologicaldifferences Synodontis 2004, 2005, 2007), as well as 2004,2005,2007),aswell fZbf]]i\j_nXk\iZiXYj etal. et al. cation (Schluter 2005; Koblmüller 2005;Koblmüller catfi 2006). Other taxa, 2006).Othertaxa, sh speciesfl (LC) is primarily aquatic but has been observed toclimb (LC)isprimarilyaquaticbuthasbeenobserved ock ock (Marijnissen that coexistinrockyshoreareas among platythelphusidspecies considerable ecologicalversatility However, thereisalso evidenceof niche divergence(Marijnissen2007). could indeedbelinkedtoecological morphological differentiationthat and catfi ( of empty stromatolite reefs,toextensivebeds cobble beaches,rockyoutcrops,and diversity ofhabitats,rangingfrom endemic fi included inthedietofnumerous prey aswellpredators.Theyare components ofthefood-webas areas. Thecrabsareimportant the speciesarefoundinrocky these habitats,butthemajorityof of mud.Crabscanbefoundinall shells, sandybeachesanddeeplayers Mastacembelus plagiostomus Lake Tanganyika offersawide sh C. stappersi

( Chrysichthys brachynema Neothauma tanganyicensis sh, suchasspinyeels etal. © Marijnissen, S.A.E.¹ ), andcatﬁ HEINZ BUSCHER ncipale, KigobeNord, 2008). 2008). sh sh ), bagrid ), bagrid

CHAPTER 6 (Synodontis dhonti), as well as cichlids and breakdown of mate recognition Lake Tanganyika crab species (Boulengerochromis microlepsis systems as water visibility is reduced (Marijnissen et al. 2008). and Lamprolus lemarii). The crabs (Donohue and Molinos 2009). Lake Tanganyika’s crabs are themselves are top invertebrate Research in Lake Tanganyika has also threatened by invasion

predators, and their relative demonstrated negative impacts of of the non-native Louisiana |

abundance can have important sedimentation on algal productivity Crayfi sh (Procambarus clarkii) and SPECIES IN THE SPOTLIGHT implications for organisms lower as well as on species diversity and the Red Claw Crayfi sh (Cherax in the food web, including algae, densities of ostracods, snails, and quadricarinatus). Both of these insect larvae, ostracods, and snails fi shes (McIntyre et al. 2004, and crayfi sh have been introduced to (Marijnissen et al. 2008, 2009). references therein). As crabs are Africa for aquaculture purposes and Lake Tanganyika is increasingly closely linked, either as predators are spreading rapidly throughout the under pressure from environmental or prey, to many other organisms continent (e.g. Arrignon et al. 1990; problems generated by human that are vulnerable to the impacts G. Howard, IUCN, pers. comm.). populations (reviewed in Lowe- of increased sedimentation, it is Considering their current presence McConnell 2003). The majority likely that they will respond by in catchments adjacent to Lake of species diversity is found shifting their foraging patterns (e.g. Tanganyika, it is likely that these in rocky near-shore areas, McIntyre et al. 2004). However, crayfi sh will move into the lake which are severely impacted by demonstrating and predicting the basin in the near future. Due to their sedimentation resulting from effects that habitat disruptions will resilience to different environmental ongoing deforestation in the lake have on the endemic crabs requires circumstances, high dispersal capacity catchment (Cohen et al. 2005). taking into account the complexity and opportunistic diet, crayfi sh Sedimentation can negatively affect of the ecosystem. Changes in the are capable of causing dramatic aquatic organisms through a range species fl ock may be delayed relative biodiversity shifts (e.g. Smart et al. of factors including deterioration of to those of other species as crabs 2002; Snyder and Evans 2006; Cruz habitat quality and heterogeneity, occupy multiple positions within and Rebelo 2007), and could cause an reduction of food intake, alteration the food web, and a considerable ecological disaster if they establish of competitive relationships, amount of overlap exists among the themselves in Lake Tanganyika. decrease of reproductive success, ecological niches of the different Without conservation effort to address environmental threats in Lake Tanganyika is the Lake Tanganyika basin, this could lead to the loss of a dynamic increasingly under pressure endemic species fl ock that provides from environmental problems unique insight into the evolution and ecological functioning of this generated by human populations unusually diverse lake system.

Platythelphusa armata (LC) is the largest and most distinctive species in the genus. Juveniles are often found sheltering within the empty shells of the gastropod Neothauma tanganyicense. © SASKIA MARIJNISSEN 197 Jg\Z`\j`ek_\jgfkc`^_k K_\

SPECIES IN THE SPOTLIGHT otamonautes raybouldi, where a tree branches. When snails. When it ﬁ nd these snails it

| the East African tree hole branching occurs it often creates a takes pieces of the snail shell back crab, is a bizarre example small hole, or bole, which fi lls with to its water-fi lled tree hole and of crab evolution. It is water. The East African tree hole masticates them, which releases Gexceptionally rare for a crab to crab has evolved to occupy this the calcium ions into solution. It live in a tree (an arboreal species), niche, adapting its behaviour and then ingests the calcium in order to CHAPTER 6 and most of those that do depend morphology accordingly. obtain suffi cient quantities to build on an external water supply – What is remarkable about this its own exoskeleton. It also eats its for example, several species of species is its natural history. This old exoskeleton when it moults. mangrove crab reside in trees for species has an apparent lack of As crabs have a hard exoskeleton protection but lay their eggs in dependence on any water body, they need to moult in order to be the sea. except that collected in tree holes able to grow and mature. Below the There are over 1,200 species as water run-off from the host tree. hard exoskeleton is soft tissue that of freshwater crab, out of which What is yet more remarkable is the expands over time. Eventually the only a handful exhibit some kind way in which it obtains the calcium hard exoskeleton will prevent any of arboreal association. The East in order to build its exoskeleton further expansion, at which stage African tree hole crab is unique (Bayliss 2002). It forages at night the process of moulting (or ecdysis) in this case. It lives solely in the or in heavy rain, when it leaves occurs. water fi lled tree holes formed its hole to search for leaf-litter It was fi rst observed and collected by Professor John Raybould in 1966, but it was not until 2004 when it was eventually described under the patronym Potamonautes raybouldi (Cumberlidge and Vannini 2004). It is a forest species found in just a few sites in east Africa, such as the East Usambara Mountains in northern Tanzania, and several sites in southern Kenya (e.g. the Shimba Hills). It has primitive lungs that enable it to spend time outside water, and elongated legs that allow it to climb up and down the tree trunks. The males have one claw that is larger than the other, which is presumably used for display and eventual mating. The females have been found protecting their young (parental care), living in the same tree holes with a brood of juvenile crabs that is presumably their own. These tree holes need Juvenile crabs in a communal tree hole. © JULIAN BAYLISS cleaning on a regular basis as leaves

198 ¹ Conservation Science Group, Department of Zoology, University of Cambridge CHAPTER 6 and other debris accumulate. The juvenile crabs are often too small to cope with moving larger leaves and debris, and would be unable

to clean the tree-hole without |

assistance from the adult female SPECIES IN THE SPOTLIGHT crab. They also need protection from predators, possibly including other crabs. The local people in the East Usambara Mountains use the water that Potamonautes raybouldi lives in as a medicinal potion. They often collect the water, which they call ‘Mazi yangodi’ (the water from holes with crabs), to give to pregnant women. The water contains calcium, which is required by pregnant women to reduce the risk of pregnancy- induced hypertension (PIH), high blood pressure (pre-eclampsia), and kidney failure, all of which can result in miscarriage. Exactly how and when the connection between the ‘crab water’ and calcium defi ciency in pregnant woman was arrived at is a mystery. The East African tree hole crab is a spectacular example of adaptation, essentially of an aquatic species not only to a terrestrial environment but also to an arboreal existence. It has lost its dependency on larger water bodies such a streams and rivers, and via the process of natural selection, through trial and error (otherwise known as evolution), it has managed to exploit a niche that crabs have rarely occupied. The further connection between the local population of the East Usambara Mountains and their use of the Mazi Yangodi further highlights this species scenario as an amazing example of natural history and a connection with a human society. However, this story of natural history is not fi nished. Exactly how the males and females fi nd each other in order to mate in closed canopy, dark forest, when each crab resides in a different tree and they are relatively short- sighted, remains an exciting mystery yet to be solved. Adult crabs defending their tree hole. © JULIAN BAYLISS 199 Chapter 7. Aquatic plants of Africa: diversity, distribution and conservation

Juffe Bignoli, D.¹

¹ Freshwater Biodiversity Unit, IUCN Species Programme, Cambridge, UK. CONTENTS 7.1 Background 202 7.2 Overview of the freshwater fl ora of mainland continental Africa 204 7.3 Conservation status 205 7.3.1 All species 205 7.3.2 Endemic species 208 7.4 Extirpated species 209 7.5 Patterns of species richness 209 7.5.1 All assessed species 209 7.5.2 Continental endemic species 211 7.5.3 Threatened species 214 7.5.4 Data Defi cient species 215 7.6 Major threats to aquatic plants 216 7.6.1 General threats 216 7.6.2 Habitat loss due to agriculture 218 7.6.3 Habitat loss due to infrastructure development 219 7.6.4 Water pollution 219 7.6.5 Water use: extraction and transfers 220 7.6.6 Invasive species 221 7.6.7 Climate change 221 7.7 Conservation recommendations 221 7.7.1 Planning and policy 221 7.7.2 Alien species removal 221 7.7.3 Catchment-scale actions 221 7.7.4 Species or site based actions 221 7.7.5 Restoration 222 7.7.6 Further research 223 7.8 Key messages 223 Species in the spotlight – A small wonder of nature saved from extinction 224 Species in the spotlight – Sailing fl owers 225 Species in the spotlight – The habitat creator 226 Species in the spotlight – Water hyacinth, a threat to the freshwater biodiversity of western Africa 227

MAIN: Freshwater habitat in the middle Zambezi river near Bovu island. © JENS KIPPING RIGHT: Myriophyllum spicatum (LCRG) is very common in the northern hemisphere and southern Africa. © SERGE MULLER RIGHT BELOW: Oxalis dines (EN) is only known from eight locations in the Western Cape, South Africa. It is found in seasonal pans in clay where it is threatened by heavy grazing and trampling. © NICK HELME 202 CHAPTER 7 | PLANTS A typical freshwater channel intheOkavango Delta. A typical freshwater groups (e.g.,White1993;Myers Africa, thefocushaspredominantly beenonterrestrial ofplantspeciesacrossbiogeographic patterns continental Although anumberofprevious studieshaveexamined Assessment 2005). nutrients, andtrappingsediments(MillenniumEcosystem ﬂ wide rangeofindirect ecosystemservicesbyreducing ﬁ sustain locallivelihoodsthrough direct use(e.g.,asfoods, life withinthesesystemsdepends.Aquaticplantshelp phytoplankton, supplytheprimaryproduction upon which to speciesinthehighertrophic levelsand,together with habitats. Theyprovide nutrientsandstructuralsupport an rivers (e.g.,CongoandZambezi).Freshwater plants play Lake Chad,andtheCameroon craterlakes),and large renowned lakes(e.g.,theGreatAfrica, Lakesineastern deltas (e.g.theNiger, Nile,andOkavangodeltas),globally ecosystems thatincludeimmensecoastalandinlandriver have ledtotheevolutionofavarietyfreshwater Africa’s hugelyvariedclimateanddiversegeomorphology 7.1 Background has collatedinformationontheconservationstatus, Mutke andBarthlott2005).Nostudyofaquaticplants ood risk,nutrientcycling,removing pollutantsandexcess bre, medicines,andstructuralmaterials),provide a

important role inthecreation andmaintenanceofthese et al. 2000;Linder2001; © JENSKIPPING aquatic plantsacross continentalmainlandAfrica. and presents asynthesisonthestatusanddistributionof compiles thefi 2009; Garcia (see Darwall of reports, eachincludingachapteronaquaticplants are availableforeachoftheregions ofAfricainaseries across thecontinent.Theresults oftheseassessments on thedistributionandstatusofaquaticfreshwater plants Here weaddress thisinformationgapbycompilingdata geographic distribution,andthreats forfreshwater species. panintheKalahariregion. A temporary et al. et al ndings from theseregional assessments . 2010;Brooks 2005;Darwall et al. et al. © JENSKIPPING 2011). Thischapter 2009;Smith et al . CHAPTER 7 | PLANTS 203 is oat on nition is nition used nition ndings from from ndings Trapa natans Trapa cult to apply to families such nition of aquatic plants as “vascular gures for the number of described gures cation scheme. Although most cation system used, potential for bias in species selection. Once it Once it selection. in species for bias potential bias apparent was no there clear that became SPF data set of species, the in the selection species, with the 353 additional was merged data set of 718 in a combined resulting fi similarities between species. The and discussed presented sets are the two data of the two data The compositions below. 7.1. All species summarised in Table sets are maps included assessments and distribution been published and are in this analysis have the IUCN Red List available through freely website (www.iucnredlist.org). be described as being Few plant families can and many apparently exclusively aquatic, somewhat drought- aquatic species are tolerant, being able to survive extended periods For this study we follow Cook’s of exposure. (1996) defi parts plants whose photosynthetically active at least, for several months permanently or, are in water or fl submerged each year, the surface of the water”. This defi particularly diffi as Compositae or Cyperaceae, which include systems which species occurring in freshwater For can also survive in temporarily dry areas. all species included in this study this reason by some authors as may not be considered true aquatics. to taxonomy arise as relating Problems no single globally accepted is currently there classifi to follow the taxonomy institutions agree by the Angiosperm Phylogeny Group proposed still many publications and are (APG) there Herbaria that use, or have used, other systems. particularly when trying problems, This creates at higher taxonomic levels studies to compare (i.e., family). For example, depending on the classifi or to the Trapaceae to belong considered As the taxonomy of plants Araceae familiy. at the time of major revisions is undergoing writing, this study follows the taxonomy applied within the IUCN Red List, which will be updated as and when a single taxonomy is agreed. Global fi aquatic vascular plants (angiosperms, gymnosperms, and ferns and allies) vary depending on the author and the defi for an aquatic plant. Cook (1996) estimated that 1 to 2% of vascular plants may be truly aquatic. On the assumption of a maximum

ed by Aquatic and Wetland Plants Aquatic and Wetland . Two data sets were analysed. First, in sets were data . Two IRIDACEAE 5 XYRIDACEAE 13 HYPOXIDACEAE 1 VIOLACEAE 1 HAEMODORACEAE 1 UMBELLIFERAE 9 GUTTIFERAE 1 TECOPHILAEACEAE 1 GRAMINEAE 15 SELAGINELLACEAE 1 GENTIANACEAE 1 SCROPHULARIACEAE 16 EUPHORBIACEAE 2 ROSACEAE 1 ERIOCAULACEAE 7 RESEDACEAE 1 ELATINACEAE 1 RANUNCULACEAE 4 DROSERACEAE 4 PRIMULACEAE 4 CYPERACEAE 129 POLYGONACEAE 3 b) Additional plant families = 353 species within 44 families b) Additional plant families = 353 species within CUCURBITACEAE 1 PODOSTEMACEAE 44 ISOETACEAE 17 CRUCIFERAE 8 PLUMBAGINACEAE 5 HYDROCHARITACEAE 35 TYPHACEAE 7 CRASSULACEAE 8 PLANTAGINACEAE 2 HALORAGACEAE 5 TRAPACEAE 1 CONVOLVULACEAE 1 PALMAE 2 COMPOSITAE 74 SALVINIACEAE 3 COMMELINACEAE 3 OXALIDACEAE 6 CERATOPHYLLACEAE 2 POTAMOGETONACEAE 17 CARYOPHYLLACEAE 4 ORCHIDACEAE 2 CABOMBACEAE 1 PONTEDERIACEAE 5 CAMPANULACEAE 6 ONAGRACEAE 3 AZOLLACEAE 2 NYMPHAEACEAE 9 CALLITRICHACEAE 4 LYTHRACEAE 7 ARACEAE 23 MENYANTHACEAE 11 BORAGINACEAE 2 LENTIBULARIACEAE 5 APONOGETONACEAE 16 MARSILEACEAE 23 AMARYLLIDACEAE 4 LEGUMINOSAE 11 ALISMATACEAE 16 LEMNACEAE 13 FamilyAMARANTHACEAE 1 Species Family LABIATAE Species 3 ACANTHACEAE 45 JUNCACEAE 40 a) Selected plant families (SPF) = 365 species within 21 families 365 species within 21 plant families (SPF) = a) Selected Family Species Family Species It was not possible to assess all described aquatic plants in Africa, so the focus was on those families identifi Cook (2004) in his publication of Southern Africa any unintended bias in the selection of to eliminate order species within families, we assessed all known aquatic species within each of 21 families. The criteria for selecting explained in Chapter 2. This was called these families are the selected plant families (SPF) data set, which includes other plant 365 species. An additional 353 species from assessed but in these cases it was not also families were leaving possible to assess all species within each family, Table 7.1. Plant families assessed and the numbers of species of and the numbers assessed families 7.1. Plant Table considered in blue are highlighted each. Families within assessed species within 65 data set = 718 aquatic. The combined to be fully families. 204 CHAPTER 7 | PLANTS regions havebeenpoorlysurveyed(e.g.CongoandCentral al. and 6,638aquaticvascularplantsglobally. Chambers Cook’s estimatemeansthatthere are between 3,319 of 315,903describedvascularplants(Chapman2009), The Makoge lakeinTanzania. fl is stillinconsistentandyettobenefi (Beentje pers.comm. described andprobably more than49,000speciesintotal diverse andfascinatingfl Continental Africaisknowntohaveoneofthemost 7.2 across mainlandcontinentalAfrica. the distributionandconservationstatusofaquaticplants provides themostcomprehensive assessmenttodateof not acompleteinventory, theinformationpresented here depending onwhichglobalestimateisused.Although 7 and18%ofallvascularaquaticplantsfoundglobally, plants assessedinthisstudytherefore accountforbetween aquatic plantismore restrictive. The484endemicaquatic aquatic macrophytes globally, althoughtheirdefi oras (BeentjeandSmith2001;Brooks (2008) mainland continentalAfrica Overview ofthefreshwater fl

estimate there tobe2,614speciesofvascular

2010 oras withmore than40,000plants . ). KnowledgeofAfrica’s plants © GITA KASTHALA. t from completed et al. ora of 2011). Some nition ofan et not, however, assumeaquaticplantswill,bydefault,be and aquaticspeciesdistributionsisnotclear. We should therefore expecttoseeasimilaritybetweenterrestrial angiosperms (Cook1996).Whetherornotweshould angiosperms haveevolvedfrom ancestorswithinterrestrial and terrestrial species,butitisacceptedthataquatic groups, withmanyfamiliesincludingbothaquatic Aquatic plantsdonottendtoformdiscrete taxonomic ofdistributionwithinAfrica. identifying patterns realms andsolacksthespatialresolution required for but theiranalysisisrestricted tolarge bio-geographic distribution ofaquaticvascularplantsattheglobalscale, al been conductedintheZambezibasin(e.g.,Timberlake et al basis forIUCN’s Africa regional southern report (Darwall Africa,andthisworkformedacentral plants ofsouthern example, Cook(2004),describedtheaquaticandwetland a fewstudiesfocusspecifi systematically describeallspecieswithinaspecifi Although acommonapproach toplantsurveysis region, are wellstudied. African Republic)whileothers,suchastheCapeFloristic 2000 . 2009).Similarly, muchworkonaquaticplantshas ). Chambers et al. (2008) reviewed thenumberand cally onaquaticplants.For cregion, et. CHAPTER 7 | PLANTS 205 , ed as likely , assessed as Least k_\_XY`kXkZi\Xkfi Prionium serratum nding given the importance of plant species to Jg\Z`\jèk_\jgfkc`^_kÆ

this volume). Figure 7.1. The proportion (%) of aquatic plant of aquatic (%) proportion 7.1. The Figure in Red List Category IUCN regional in each species continental Africa. mainland to its global Red List Status. Those species correspond or with less than 5% of their not native to the region, not included within global distribution within Africa, were for all species results the analyses. Section 7.3.1 presents for Section 7.3.2 shows the results assessed, whereas Africa. only those species endemic to continental 7.3.1 All species assessed as Least of all species were Fifty eight percent common 7.2), being relatively Concern 7.1 and Table (Figure identiﬁ with no major threats and widespread to impact their overall population viability. This represents This represents to impact their overall population viability. a positive ﬁ ecosystems. Some the function and stability of freshwater species, such as Concern, noted for their importance as ecosystem are new habitat for many other species engineers creating (see Number endemics of regional of regional

et al. et al. species Number of 1994). Myers et al. 1994; Linder 2001). ed include: the mountain ranges of ed include: the mountain ranges 2005; Beentje Extinct In The Wild 1 1 Critically Endangered (CR)Critically Endangered (EN)Endangered (VU)Vulnerable (NT)Near Threatened 36Least Concern (LC)cient (DD) Data Defi 31 32 57 78 415 100 23 41 69 238 80 IUCN Regional Red List Category uctuations (changes in climate throughout in climate throughout uctuations (changes et al. et al. Conservation Status Total 718 484 Threatened Threatened Categories effectively captured within conservation plans based plans conservation within captured effectively distributions. species of terrestrial on knowledge purely what is to examine it is a useful exercise Nevertheless, many studies are There species. terrestrial known for plant on terrestrial we can extract information which from Africa patternsdistribution types across and vegetation Linder 2001; Kuper 1983; White 1993; (e.g., White 2004; Kier 7.3 conducted were Assessments of species extinction risk following the IUCN Guidelines on application of the IUCN Red List Categories and Criteria at national or regional Red List status levels (IUCN 2003), noting that the regional of any species endemic to mainland continental Africa will IUCN Red List Category, Numbers of species in each regional 7.2. Table as applied to mainland continental Africa. (2000) also used plant distributions as a key component (2000) also used plant global Biodiversity for delineating 21 within the process studied patternsHotspots. Linder (2001) diversity of plant a total of 1,818 species in sub- and endemism for that, while overall Africa and concluded Saharan tropical to precipitation related species richness patterns are to closely related more levels, patterns of endemism are paleoclimatic fl of species evolutionary history). Some examples Africa’s identifi rich areas (High Atlas and Middle Atlas Algeria and Tunisia Morocco, to Ghana), mountain ranges); Upper Guinea (Sierra Leone Equatorial Guinea Lower Guinea (Nigeria, Cameroon, and (Tanzania and Gabon); the East African coastal strip the Lake Kivu Kenya); the Zambezi-Congo catchment; catchment in easternCongo, Rwanda and Burundi; the easternof Zimbabwe and the adjacent highlands Floristic Region. mountains of Mozambique; and the Cape of endemism include the High Commonly cited centres Upper Guinea Atlas and Middle Atlas mountain ranges; catchment; the and Lower Guinea; the Zambezi-Congo Cape East African coastal strip; the Lake Kivu catchment; Floristic Region; and Huilla in southern Angola (White 1993; Beentje 206 CHAPTER 7 | PLANTS © JEAN-PAUL GHOGUE restricted distribution and threats fromwaterpollution. distribution andthreats restricted provinces. Ithas beenassessedasVulnerable duetoits ofthesouth-west toriversinthecoastalarea is restricted Macropodiella pellucida are considered toberegionally threatened. threatened. Africa Bycontrast,24%ofspeciesinnorthern Africa, forexample,only5%ofspeciesare considered large regional differences inthelevelofthreat. Insouthern IUCN RedListthreatened categories(Table 7.2),with One infi Red ListCategoryinmainlandcontinentalAfrica. Figure 7.2.Theproportion (%)ofaquaticplantspecieswithinthefullyassessedfamiliesineachregional IUCN ve species(20%)are assignedtooneofthe (VU)isendemictoCameroonand

existing dam. byan itisthreatened is knownfromonly onesitewhere Winklerella dichotoma this highlevelofthreat isarefl threatened (Vie IUCN RedListCategoriesandCriteria.Ofthese,70%are 3% ofknownplantspecieshavebeenassessedusingthe of thetotalnativefl (2000) other fi In theglobalcontextthislevelofthreat isconsistentwith

concluded that72%ofendemicspeciesand19% ndings. InEcuador, forexample,Valencia © JEAN-PAUL GHOGUE et al. ora are threatened. Globally, lessthan 2008), but it has been suggested that 2008),butithasbeensuggestedthat (CR)isaCameroon endemic.It ection ofbiasamongstthe et al. et al. CHAPTER 7 | PLANTS 207

et ow et al. (VU). Macropodiella with 27 of the

, for example, has (CR), cations to the river’s fl cations to the river’s Ledermanniella keayi Ledermanniella kamerunensis Winklerella dichotoma cient information to assess the conservation (EN), and cient (DD). The DD category is assigned when 2008), requiring seasonal inundation, clean water, and and seasonal inundation, clean water, 2008), requiring Although all aquatic species within the family Although all aquatic species within it has the Podostemaceae have not yet been assessed, species highest number of threatened With 86 species of 44 assessed species threatened. Podostemaceae in continental Africa (Rutishauser within this family 2007), this means at least 31% of species at risk of extinction. Podostemaceae is the largest are (Chambers exclusively aquatic family of angiosperms al most of the high light intensity (Philbrick 2004). Globally, endemic to small geographic areas. Podostemaceae are and to the centre restricted are Africa, species Across west of the continent and often have a limited distribution occurring in a single cataract or river (Cook 1996). Such localised distributions make these species more vulnerable to habitat degradation and alterations to river by water abstraction, impoundment caused hydrology and sedimentation. only been found at a single site on Mount Oku and the Ijim It has been listed as Critically in Cameroon. Ridge area of habitat degradation associated result as a Endangered with water abstraction, modifi agricultural development. Other and upstream regime, species in this range threatened examples of restricted family include listed as of all species assessed are Fourteen percent Data Defi is insuffi there status of the species and, as such, does not indicate that pellucida . 2008). A © JEAN-PAUL GHOGUE © JEAN-PAUL et al spp.). Finally, the Isoetaceae, spp.). Finally, (LC) is endemic to central Africa. This plant is very (LC) is endemic to central Africa. This plant specialized in term found in of ecology as it is only (VU) is a threatened species only (VU) is a threatened cant differences in the numbers of in the cant differences Nymphoides expected to be threatened (Vie (Vie expected to be threatened a priori owing waters such as rapids and waterfalls. threatened species within those plant families where species within those plant families where threatened assessed. High numbers all aquatic species were found within the families are species of threatened Aponogetonaceae, Menyanthaceae, and Isoetaceae is a family of fully aquatic 7.2). Aponogetonaceae (Figure species, mainly endemic to Africa, found in seasonal and valued locally temporal marshes and pools. They are for food and for decorative purposes. As an example, Aponogeton azureus locality in northern a single known from Namibia, where to harvesting and grazing pressures. due it is threatened found in still waters and Species of Menyanthaceae are their use as have an important economic value through ornamental species, mainly in aquariums and water (e.g. gardens botanical community to focus primarily on species thatbotanical community to focus primarily were Kew, carried out by Royal Botanical Gardens report recent IUCN, usingthe Natural History Museum of London, and balanced picture a more provides a sampled approach, (for globally threatened concluding that 22% of plants are information see RBG 2010). more signifi are There submerged, to as quillworts, can grow commonly referred basis of a On the or seasonally submerged. emergent by considered quillworts are number of unusual features, of ancient trees some botanists to be the last remnants Some species of quillwort are the early Triassic. from useful indicators of declining water quality being highly intolerant to nutrient enrichment (Cook 2004). Dicraeanthus africanus fast fl 208 CHAPTER 7 | PLANTS Category inmainlandcontinentalAfrica. plant speciesineachregional IUCNRedList Figure 7.3.Theproportion (%)ofendemicaquatic from taxonomicuncertainty. for whichthedistributionrangeisunclearorsuffer resolved. MostoftheplantslistedasDDare rare species prove tobethreatened whenthesegapsinknowledgeare the speciesisnotatrisk.Manyofthesemay Cape-pondweed. ofWaterblommetjiebredie,widely usedforhumanconsumption.Forexample,therecipe localdish,includes arenowned Aponogeton dystachyos © CRAIGHILTON-TAYLOR (LC),knownasCape-pondweedisendemictoSouthAfrica.Thebudsandfl are justafew examples of in Cameroon) are alllisted asCriticallyEndangered, and and Congo), to SouthAfrica), extinct. Speciessuchas plants nativetoAfricaare atriskofbecomingglobally assessed (20%).Thismeans that oneinfouraquatic endemic species(26%)ishigherthanthatforall Date Defi categories (CR,EN,VU),and17%were considered while aquarterfallintooneofthethreatenedConcern, Of theendemicspeciesassessed,49%are listedasLeast traditions (seeSection7.6.). uniqueness andinsomecasesare linkedtoregional have focusedonendemicspeciesthatrepresent local Africa. Historically, mostconservationefforts andstudies Lagarosiphon cordofanus Kenya.Others,suchas one seasonalpoolinsouthern nigroreticulata being confi risk ofextinction.Somehaveveryrestricted ranges,often risk ofextinctionwithinAfricaisequivalenttotheirglobal to reiterate thatinthecaseoftheseendemicspecies,their are endemictomainlandcontinentalAfrica.Itisimportant Four hundred andeightyfourofthe718speciesassessed 7.3.2 Endemicspecies Ledermanniella batangesis Ledermanniella cient (Figure 7.3).Theproportion ofthreatened Cyperus microumbellatus ned toasinglearea orsite,suchas (CR) , Drosera insolita which iscurrently onlyknownfrom (LC), are widespread throughout

very restricted rangespecies in Cadiscus aquaticus (endemicto owers ofthisplantare (endemictoKenya), (endemictoD.R.

a singlesite (endemic Isoetes

CHAPTER 7 | PLANTS 209 cient species, distribution cient species, c. With this limitation in mind, c. With this limitation in mind, rms that little, if any, bias is introduced bias is introduced rms that little, if any, ed. The only notable difference between the ed. The only notable difference data were unavailable at the catchment scale, and in catchment scale, unavailable at the data were to country boundaries. mapped species were these case of spatial data is the course resolution This relatively could be obtained for countries best information that data are Congo, and Congo where such as Angola, D. R. and unspecifi often scarce the most provides below presented the spatial overview of aquatic to date of the distribution complete picture It is, however, continental Africa. plant species throughout the to close is required research also clear that further species. information gap for many 7.5.1 All assessed species Species richness patterns for species within the selected 7.4a) and the complete data plants families (SPF) (Figure minimal 7.4b) reveal set for all assessed species (Figure between the overall patterns of species differences richness. This confi families for which not inclusion of species from through and assessed. all aquatic species have yet been mapped therefore The pooling of all species within the analysis seems justifi ranges adjacent two richness maps lies in the mountain 7.5 Patterns richness species of to river possible where mapped are Species distributions in particular for widespread However, sub-catchments. Least Concern and Data Defi © WILLIAM DARWALL rmed Ottelia ed and and ed 8jdXcc , this volume. (Nymphaceae) (EW) cient or Critically conservation efforts – its conservation efforts ex-situ Jg\Z`\jèk_\jgfkc`^_kÆ

Nymphaea thermarum (Hydrocharitaceae) has only been recorded been recorded has only (Hydrocharitaceae) cient species are a priority for further research, for further research, a priority species are cient ciency, the true proportion of threatened species of threatened the true proportion ciency, rm if the species is at risk of extinction or whether the species is at risk of extinction or whether rm if nfe[\if]eXkli\jXm\[]ifd\okèZk`fe need of targeted conservation actions if they are to survive to survive if they are actions conservation of targeted need long-term. in the Data Defi can be identifi found to be threatened as those later action taken to ensure their survival. For example, survival. For example, their to ensure action taken brachyphylla on information is no reliable there once in Sudan and will or its distribution. Further research threats current confi exist. With such high levels of other sub-populations data defi later found to be all DD species are could rise to 40% if threatened. Nymphaea and Nymphoides spp. in the Okavango Delta. now only survives due to story is given in 7.4 Extirpated species (or Regionally For a species to be assessed as Extinct doubt that the last must be no reasonable Extinct) there be confi individual has died. Until extinction can species may be listed Data Defi this into account, (Possibly Extinct). Taking Endangered the selected families has been only one aquatic plant from in a single listed as Extinct in the Wild. Originally found site in Rwanda, 210 CHAPTER 7 | PLANTS © WILLIAM DARWALL throughout manycountriesacrossAfrica. widespread Trapa Natans families (SPF);andb)allspecies.Speciesrichness=numberofspeciesperriver/lakesub-catchment. Figure 7.4.Thedistributionofaquaticplantspecies across mainlandcontinentalAfricafor:a)Selectedplant (LC),alsoknown aswaterchestnut,is extending from MountCameroon equatorial to thewestern The highestdensity ofspeciesisseenwithin anarea for plantspeciesrichness. argues thatprecipitationprovide patterns agoodindicator hasbeenpointedoutbyLinder (2001),who similar pattern Africa inparticular.in theLowerGuineaandeastern A across thetropics, withhighnumbersofspeciesobserved The greatest densityofspeciesfollowsabroad band levels ofendemism. the Kivucentre duetoitshighspeciesdiversityand 7.4b). Thisarea hasbeenhighlightedbyLinder(2001)as that isonlyrevealed whenallspeciesare pooled(Figure and Rwanda,whichshowanarea ofsignifi D.R.Congo,includingBurundi to theriftlakesinwestern cant richness cant CHAPTER 7 | PLANTS 211 ora ects 2000), et al. ciency rather than a true biogeographic pattern.ciency rather Mediterranean Biodiversity Hotspot (Myers of Eurasian and African fl the intersection where and fauna has created an area of exceptionally high an area and fauna has created biodiversity. which includes Species richness in the Upper Guinea area, Liberia, Sierra Leone, Guinea, and Congo, and northern with D. R. Congo and Angola, especially on the borders Zambia, was lower than expected. This most likely refl data defi 7.5.2 Continental endemic species as for overall species endemism are, of species Centres richness, in the southern Gulf of Guinea drainages (Mount highlands) and the Lower westernCameroon, Cameroon crater lakes, where individual sub-catchments are recorded recorded individual sub-catchments are crater lakes, where of high to contain up to 175 species. Other notable regions species richness include the Lower Niger Delta in Nigeria, Equatorial Guinea, Gabon and Congo. In eastern Africa the great richness extends across of high species an area the Lake Bangweulu rift lakes and river systems. Finally, basin, Lake Mweru basin and southern D. R. Congo are also were also species rich. A number of these regions highlighted by Linder (2001). the Cape Although less diverse than the equatorial regions, Africa, KwaZulu-Natal, Mpumalanga and in South fold area Limpopo, also support notably high numbers of species. In northern Africa, the Atlantic and Mediterranean coasts and the Nile represent and Tunisia, Algeria of Morocco, of biodiversity known as the centre a globally renowned Cadiscus aquaticus (CR) is endemic to the Western Cape between Groenekloof and Saldanha Bay. It is restricted to vernal pools where it is threatened through overgrazing by livestock, infi lling of wetlands, invasion by alien grasses and eutrophication. © NICK HELME

Cotula myriophylloides (CR) is endemic to the Cape Peninsula in the Western Cape, South Africa. This species had not been collected for the past 30 years and was thought possibly extinct until this population was discovered at Kleinmond in October 2008. Its main threats are urbanisation and invasive alien species. © NICK HELME CHAPTER 7 | PLANTS 213 Pulicaria the orchid the orchid

(CR), (CR), and (CR), all of which are threatened by threatened (CR), all of which are Juncus maroccanus Epilobium numidicum (CR), laginoides the High Atlas and Middle Atlas mountain ranges, and within the upper Nile Delta. Examples of species endemic include to the area ﬁ habitat loss and fragmentation. This is a common threat an unfortunate to species in northern Africa, representing development that has left less than of human by-product intact today. 5% of the hotspot area of species endemism extends Another important area the southern across of Lake Victoria north-east from lakes of Kenya and the Eastern Rift to Lake Tanganyika. with part of the Somali-Massai and coincides This area of plant endemism also described by centres Afromontane White (1993). The Okavango delta, highlands of Lesotho, Serapias stenopetala 2000). ed centres et al. are found in this area, area, found in this are

Macropodiella pellucida (VU) (VU), Saxicolella laciniata Ledermanniella cristata Niger Delta (Figure 7.5). Species of Podostemaceae, such Niger Delta (Figure as (EN) and Figure 7.5. The distribution of aquatic plant species endemic to mainland continental Africa. Africa. continental to mainland endemic species plant of aquatic distribution 7.5. The Figure sub-catchment. per river/lake of species = number richness Species where they face threats from tourism and water pollution. tourism and water pollution. from they face threats where The overall pattern with that reported of endemism agrees by Linder (2001) and White (1993), who identifi well with those shown in of endemism corresponding the Lower and Upper of these are 7.5. Examples Figure the Cape Floristic Region (Linder 2001) Guinea Centres, and the Mediterranean and the Guineo-Congolian Regional (White 1983). of endemism Centres As mentioned above, northern Africa is an important of endemism (White 1993) and is part of centre regional the Mediterranean Biodiversity Hotspot (Myers found across High numbers of endemic aquatic plants are 214 CHAPTER 7 | PLANTS richness =numberofspeciesperriver/lakesub-catchment. Figure 7.6.Thedistributionofthreatened speciesofaquaticplantsacross mainlandcontinentalAfrica.Species richness andendemism. Highnumbers ofthreatened coincide withthose already identifi Areas withhighestnumbersofthreatened specieslargely 7.5.3 Threatened species available datainthiscurrent project. centre ofendemism, butthiswasnotconfi identifi of thelackinformationfortheseareas. Linderalso system andsouthofLakeVictoria maybeanartefact Apparent lownumbersofspeciesintheCongoRiver Congo alsoholdimportantnumbersofendemicspecies. and theborder zoneofnorthAngolaandsouth-westD.R. ed theCongo-Zambeziwatershedasaregional ed as centres ofspecies rmed bythe intense (Rhazi andGrillas 2010).These include thecoastal wetlands foragriculture andcoastaldevelopment are most species are foundinthoseareas where drainage of Africathehighestnumbers ofthreatenedIn northern both ofwhichare restricted totheseareas. Acanthus latisepalus prevalent. Suchimpacts areforspeciessuchas ofconcern changes infl aquatic plants,suchasdeforestation leadingtosiltationand areas inthisregion, majorthreats tofreshwater systemsand ecoregions (Figure 7.6).Althoughthere are manyprotected Ogooue-Nyanga-Kouilou-Niari andLowerNiger-Beune Mount Cameroon,GulfofGuineadrainages,the southern species areequatorialcraterlakes, foundinthewestern ow regimes, andpollutionfrom miningare still (VU)and Dicraeanthus zehnderi (CR), (CR), CHAPTER 7 | PLANTS 215 Oxalis cient (see cient (VU), (CR). Aponogeton angustifolius cient (DD) species are located in the cient (DD) species are Cadiscus aquaticus ecting the often-poor state of knowledge cient species cient cant numbers of DD species include the lower (CR) and Cape Peninsula. This is an area where agriculture, urbanization agriculture, where Cape Peninsula. This is an area many aquatic species threatening and invasive species are holds many endemic and(Sieben 2009). The Cape Fold area as species such rare natans 7.5.4 Data Defi assessed as Data Defi are of species A proportion section 3.1) refl to aquatic plants (Figure on the distribution and threats 7.7). Most Data Defi northern of endemism extending Upper Guinea centre and Gambia River basins. Other areas the Senegal across with signifi (CR) is (EN), for eri Africa. continental mainland across plants of aquatic cient species laginoides Ethulia scheffl Pulicaria fi plain and the Rif, Tell Atlas and Middle Atlas mountain plain and the Rif, Tell and Algeria, and the Mediterranean ranges in Morocco and Algeria. coast of Tunisia location on the Mediterranean coast one only known from by urbanization. it is threatened of Algeria, where In eastern species occur in south- Africa many threatened infrastructure where east Kenya, especially near Nairobi, pose important threats development and agriculture habitats. plant to freshwater example, is endemic to eastern Africa and is threatened near Nairobi. land conversion by severe In southern is an important concentration of Africa there especially in the in the Cape Fold area, species threatened Figure 7.7. The distribution of Data Defi of Data distribution 7.7. The Figure due be mapped could not all species Note: sub-catchment. per river/lake of species = number richness Species geographic ranges. information on their to lack of 216 CHAPTER 7 | PLANTS for water, contributed tothepollution ofsurface andground industry during the lastdecadehasincreased thedemand Globally, expansion ofagriculture and themanufacturing waters ofAfricaisthelossand degradationofhabitat. The mostsignifi 7.6 Majorthreats toaquaticplants analysis. Notethatmanyspecieshavemore than onethreat listed. continental Africa.Onlythosespeciesintheplantfamiliescomprehensively assessed(SFP)are includedinthe Figure 7.8.Percentages ofaquaticplantspeciesaffected bythemajorcategoriesofthreat across mainland taxonomic uncertainty. species. Inthesebetterstudiedareas themainissueis well surveyed,alsocontainrelatively highnumbersofDD other areas suchastheCapeFold,whichare relatively Mediterranean coastofMorocco, AlgeriaandTunisia, attributed tolowsamplingintensityinareas suchasthe Although thehighconcentrationofDDspeciescanbe information onmanyaquaticspeciesislimited. Victoria riftare basinandthenorth-eastern areas forwhich Africa,theLake of endemismlies(White1993).Ineastern the countrywhere theso-calledZambezianregional centre numbers ofDDspecies,particularlyinthesouthpart region withAngolaandZambia.itselfhassignifi a highnumberofDDspecies,particularlyalongtheborder Ogun andOshunriversinNigeria.D.R.Congoalsocontains incorporates importantfreshwater systemssuchasthe Bright Drainagesecoregion (Thieme and upperNigerDelta,theLakeChaddrainage, cant threat toaquaticspeciesinthe inland et al. 2005), which 2005),which cant cant reports (seeDarwall at aregional scaleisprovided through aseriesofregional but more detailedinformationonthreats to aquaticplants be detectedthrough thissynthesisatthecontinentalscale Due tomattersofscale,somelocalisedthreats maynot plants, are dealtwithinmore detailbelow. the manyotheractivitieswhichthreaten Africa’s aquatic subject toeutrophication (Nyenje the highproportion (28%)oflakesandreservoirs inAfrica and probably represents asignifi remains evenlower(around 30%insub-SaharanAfrica), sanitation (UNDESA2010),thelevelofsewagetreatment Africa isservedwithsafedrinkingwaterandadequate Assessment 2005).Whileonly60%ofthepopulationin habitats, andAfricaisnoexception(MillenniumEcosystem water, andhasdegradedordestroyed manywetland over the period1990to 2000.Furthermore, an additional Globally, Africahas seenthelargest regional population rise 7.6.1 Generalthreats subject toaparticularthreat. percentage valuesrepresent theproportion ofallspecies 7.8. Resultspresented are absolutevalueswhere the The majorthreats toaquaticplantsare shown inFigure et al. 2009;Garcia et al. et al. 2010;Brooks 2005;Darwall cant factorcontributingto et al et al. . 2009). These, and . 2009).These,and et al. 2011). 2009;Smith CHAPTER 7 | PLANTS 217 ) grows in poorly RG (EN © D. TWEDDLE © SERGE MULLER Marsilea strigosa mineralized temporary pools of Tunisia, Algeria and mineralized temporary pools of Tunisia, Morocco. It faces a number of threats, such as overgrazing, water extraction, and habitat loss due to changes in land- use related to agriculture and the construction of road infrastructures. The clover fern For threatened species alone (Figure 7.8) habitat loss and species alone (Figure For threatened also the major causes of species decline, pollution are by habitat loss due to with 34% of species affected and 36% by habitat loss due to infrastructure agriculture to 25% of pollution is a major threat development. Water species. threatened cient information ed as the second largest the second largest ed as ed as the other major causes of uents. Finally, drought and invasive alien drought uents. Finally, cient (DD) may also be impacted by threats by threats cient (DD) may also be impacted 65 percent increase in the population is expected between increase 65 percent the population 2000 and 2025 (WHO/UNICEF 2000). As of urban areas, rises, this is closely followed by expansion putting ever- industry and infrastructure, agriculture, on wetland ecosystems. The extent pressure increasing examined below. and impacts of such developments are not were Although 58% of the species assessed to be at risk of extinction, this does not mean considered Species listed as Least Concern they face no threats. (LC) or Data Defi which, if not halted or minimized, could lead to the species DD species may already in the future. becoming threatened lack suffi we currently but be threatened, and for all LC recorded to make this assessment. Threats included in the analysis. therefore DD species are to aquatic Habitat loss or degradation is the major threat plants, impacting 26% of the species assessed. The main cause of habitat destruction is agricultural expansion through draining wetland of land (through irrigation, reclamation development, farming. Infrastructure and livestock areas), such as urban expansion, industrial development and dam identifi construction, are habitat loss. Pollution is identifi 15% of the species assessed. This is affecting major threat, mostly attributed to water pollution by agricultural, industrial and domestic effl species impact 8 and 5% of species, respectively. The extensive Mweke swamp at Kapikili in the Upper Zambezi catchment in Zambia. The extensive Mweke swamp at Kapikili in the Figure 7.9. Numbers of aquatic plant species associated with each of the main habitat types across mainland continental Africa. PLANTS

| CHAPTER 7

The numbers of species recorded against each major habitat type are presented in Figure 7.9. The chart illustrates that the highest numbers of threatened species are found in habitats such as permanent rivers, creeks and waterfalls, followed by seasonal and intermittent marshes and pools. Permanent rivers, especially rapid waters and waterfalls, are a potential source of hydropower, water for agriculture, urban and industrial use, and provide many other important resources such as food ﬁ sh, ﬁ bre, and building materials. Consequently, despite their great value in terms of the ecosystem services provided, they are more often subject to the pressures associated with development than are other habitats. Intermittent and seasonal pools and marshes are at risk from drainage or conversion for associated agricultural activities.

7.6.2 Habitat loss due to agriculture

Agricultural expansion is responsible not only for the loss of physical space for species to grow but also the Pilularia minuta (CRRG) is a small amphibious fern found in degradation of many habitats. Conversion (clearing or Algeria, Morocco and Tunisia. Its population is in decline in transformation) or drainage for agricultural development northern Africa as a result of its very small and fragmented have been the principal causes of inland wetland loss area of distribution, combined with anthropogenic threats to worldwide (Millennium Ecosystem Assessment 2005). its habitat of temporary pools. © SERGE MULLER Across Africa commercial farmers drain wetlands for planting crops such as Amadumbe (Colocasia esculenta) or sugar cane (Saccharum offi cinale). Subsistence farmers to help African people produce the food they need. One may also pose a threat to species and their habitats where of the most important initiatives of the past years is The they are restricted to limited areas (Sieben 2009). Alliance for a Green Revolution in Africa (AGRA). AGRA is an African-led partnership whose main goal is “…to As the population rises in Africa it becomes increasingly achieve a food secure and prosperous Africa through evident that the intensifi cation and effi ciency of agricultural the promotion of rapid, sustainable agricultural growth production has to increase too. The main challenge is to based on smallholder farmers” (AGRA 2010). AGRA’s combine intensifi cation with conservation of the underlying strategy includes increasing two- or three-fold crop environmental services provided by wetlands. There are yields to address food shortages. This will be achieved 218 many initiatives that seek to increase productivity in order through improving soil fertility using organic and inorganic CHAPTER 7 | PLANTS 219

habitat . 2009). The main Juncus maroccanus ed, with only a few Acanthus latisepalus et al (VU), ora, eutrophication leads to ora, eutrophication (EN) are all threatened by all threatened (EN) are Isoetes capensis 2010). Species such as

Aponogeton angustifolius (VU), suitable habitats for many plant species (Rhazi and Grillas suitable habitats for many plant species (Rhazi is a areas 2010). In South Africa urbanization in coastal for many species, as it not only reduces serious threat pollution in of suitable habitat but also increases areas infrastructure adjacent wetlands (Sieben 2009). Increasing construction of development, which includes urbanization, has also been described as dams, or irrigation structures, to aquatic plants in central Africa one of the major threats (Ghogue (CR) and development. infrastructure pollution 7.6.4 Water a serious ongoing problem presents Eutrophication for African water bodies (Nyenje (N) and input of mainly nitrogen cause is an increased in phosphorus (P) into the aquatic system resulting “blooms” of phytoplankton or other plant species that are able to quickly utilize the excess nutrients. Eutrophication changes in species species mortality, leads to increased In assemblages, and an overall loss of aquatic diversity. the case of native aquatic fl plant assemblages becoming simplifi commonly at a cost to dominant species remaining, (VU) is restricted to Cape Province in South Africa where it is threatened by habitat loss due to © NICK HELME fertilizer, using “high quality local adapted varieties” using “high quality local fertilizer, and research increasing programmes, breeding through policies that provide capacity building, and promoting support to farmers (AGRA 2010). comprehensive to be are If initiatives such as that mentioned above sustainability must be included successful, environmental Services as a key component within planning processes. in turn ecosystems, which are by wetland provided dependent upon aquatic plants as the primary producers and habitat builders, underpin the success of these future development ambitions. Reliable information on aquatic study, this current through species, such as provided be fully integrated within these initiatives must therefore in an as pledged, they move forward, to ensure if we are Without the effective way. sustainable environmentally integration of such information, wetlands and their wildlife may be impacted in ways that might have been avoided if the necessary information had been accessed. development 7.6.3 Habitat loss due to infrastructure development have been infrastructure Impacts from the whole continent. In northern throughout recorded to tourism in coastal areas Africa development related availability of water and affecting high, is extremely Aponogeton angustifolius urbanization. 220 CHAPTER 7 | PLANTS A waterbodyinMoroccoaffectedbyover-abstraction ofwater. water quality andspecies survival.For example, accumulate inrivers andotherfreshwater bodies,affecting and organic wastes from leakingsewage systemscan mining activities(suchaslead,cadmium, iron, andcopper) survival andwaterqualityinwetland areas. Pollutantsfrom (e.g., thetsetsefl pesticides usedinagriculture andagainsthumanpestvectors been recorded asthreats toplantspecies.Forexample, species, pollutantsfrom arangeofothersources havealso Although eutrophication hasamajorimpactonplant the riftlakesinEthiopia(Nyenje Zimbabwe, LakeAlbert,theZeekoevleiinSouthAfrica,and are numerous andincludeLakeVictoria, LakeChivero in Examples ofAfricanlakesaffected byeutrophication eventually discharging wastewaterintoriverandlakes. remainder isdisposedofviaonsitesanitationsystems, than 30%istreated insewagetreatment plants, whilethe produced bycitiesinsub-SaharanAfrica,atpresent, less or miningactivities.Ofthetotalamountsofwastewater eutrophication maycomefrom wastewater, pesticides, buffering capacityofthewetland.Pollutantsthatcause in urbanareas, haveincreased nutrientloadingbeyondthe freshwater habitats,andincreased wastewaterproduction However, inmanycasestherapidratesofencroachment on (N andP)cantherefore actasabuffer toeutrophication. plants infreshwater systemsisabletoretain excessnutrients reproduction strategies.Ahealthycommunityofaquatic species thatcantakeadvantageoftheirmore competitive eutrophication mayfavourtheproliferation ofinvasive specialists andendemicspecies(Sieben2009).Inaddition, y) are anemerging problem forspecies et al. 2009). Limonium Limonium © JP-BOUDOT As discussed above,irrigation ispredicted toincrease irrigation followed bydomesticuseandindustrial use. By sector, thehighestwateruseis for agriculture through and theirvectors. fi as gravelspawningbedsforfi another withthesubsequentloss ofbenthichabitats(such water, introduction offi changes inthephysicalandchemicalstatusofriver’s rivers andincreased fl specifi functioning, leadingtoafrequent lossofbiodiversity. More with majorimplicationsforchannelintegrityandecological water-rich catchmentstothosewhere waterislimited, 1998). Thishasledtoanincrease inwatertransfersfrom Nile, Sanga,Chari-LogoneandVolta (DonkorandWolde river basins–theCongo,Niger, Ogooue(Gabon),Zambezi, of totalwaterresources concentratedwithineightmajor continent containedwithintheCongoRiverbasin,and75% continent, withabout50%ofthetotalsurfacewateron Water resources are unevenlydistributedacross theAfrican 7.6.5 Water use:extractionandtransfers (VU). (CR), agricultural orminingactivitiesinclude agriculture. Otherspeciesaffected bywaterpollutiondueto and Morocco thatisdirectly impactedbypesticidesusedin duriaei sh species,fl cally, watertransfersleadtofl Aponogeton bogneri (Plumbaginaceae) (CR) is a plant native to Algeria (Plumbaginaceae) (CR)isaplantnativetoAlgeria oating aquaticplants,andanimaldiseases ow intherecipient rivers,alongwith ne sedimentsfrom oneriver to (EN),and shes), andthespread ofalien ow reduction indonor Cadiscus aquaticus Drosera bequaertii

CHAPTER 7 | PLANTS 221 2008). The importance (also see Chapter 9,

. (also see Chapter 9, et al ting human well-being and conserving cally to “…identify the most important ). rst option is expensive and time consuming 2005 et al. Catchment-scale actions this volume) Species or site based actions this volume) 7.7.3 to be effective Although site based conservation has proved species in threatened a key tool for protecting and remains the best way tothe short and medium term, in many cases is to combine both catchment achieve conservation targets (Boyd and site scale approaches biodiversity of riparian ecological corridors for maintaining and natural patterns known (Forman 1995; Bunn and is well Arthington 2002; Bennet 2003). comes from One example of the landscape scale approach This vastongoing work in the Miombo-Mupane woodlands. of savanna is irrigated by rivers, pools and pans across area Angola, Zambia, Namibia, Botswana and Zimbabwe. It is a highly dependent local communities are where diverse area conservation Several large-scale resources. on water-related initiatives seek to manage healthy ecosystems within this ecosystem while benefi International the Conservation nature. is working towards this unique establishment of ecological corridors across landscape, specifi within Miombo-Mopane, to describe and areas freshwater the key factors driving and maintaining understand what are that that will ensure such systems, and to develop programs systems provide the essential services that these freshwater (Conservation International the future” viable into will remain 2010). 7.7.4 Site based actions also have their place within the portfolio of options available to managers. The Alliance for Zero methods as Environmental Flows and Integrated River Basin Integrated Flows and as Environmental methods as many often required, is an approach Such Management. invasive as pollution and to aquatic plants, such threats catchment, a throughout rapidly spread species, will source the initial from species sometime distant impacting of the threat. 7.7.2 Alien species removal eradicate invasive species and to control measures Effective to enable native species to successfully needed if we are are by non-native former habitats taken over re-colonize invasive the only way of controlling species. In many cases or chemical plants is periodic physical removal freshwater The fi control. species and be harmful for non-target and the second may is biological control, option A third overall ecosystem health. use success through has been some reported for which there this techniqueof plant feeding insects and pathogens, but consideration and needs careful is not completely risk free (Moran cally water Azolla , Salvinia

% Pistia striatiodes ( are well-known invasive are 2005). The issues around 2005). The issues around (water hyacinth), NXk\i_pXZ`ek_#Xk_i\Xkkf et al. ed important sites, the most cient use of existing irrigated areas areas use of existing irrigated cient Typha australis, Azolla pinnata Typha Ludwigia stolonifera . 2006). Whatever the outcome, an Integrated the outcome, . 2006). Whatever be that could process in the production ciencies or Eichornia crassipes (kariba weed), et al , ) Jg\Z`\j`ek_\jgfkc`^_kÆ

liculiodes 7.6.7 Climate change 8% of the species to affect reported are Although droughts assessed, the impact of climate change was not specifi k_\]i\j_nXk\iY`f[`m\ij`kpf]n\jk\ie8]i`ZX considered as a threat. The potential impacts of climate threat. as a considered Africa are, species throughout change to freshwater discussed in a dedicated section within Chapter however, 8, this volume. lettuce molesta enormously in the future in response to growing demands demands growing to in response future in the enormously to the need However, agriculture. productive more for a has been immediate solution as the areas expand irrigated reported through region, for the sub-Saharan challenged ongoing defi effi more solved through strategy is necessary if we Management Resource Water water while increasing biodiversity freshwater to protect are use for irrigation. 7.6.6 Invasive species to native pose an important threat Alien invasive plants or competition for resources direct species, either through changing vegetation dynamics ultimately indirectly through or loss of plant diversity (seeleading to habitat degradation also Chapter 8). Species such as fi examined in of water hyacinth are and control the spread (Riddell species. Their harm to native habitats has been widely species. Their harm to native habitats has has their control documented, and the cost associated with of water hyacinth to be high. For example, the control proven Africa. to many wetlands throughout is an important problem annual cost ofAlimini and Akinyeminju (1990) calculated the USD from clearing 1km2 of water hyacinth in Nigeria ranged to USD 4,400 for chemical control. 9,500 for manual control environmental Calculated costs will be even higher if the wetlands from costs of using chemicals to eradicate weeds as invasive in hyacinth has been reported included. Water are and the Middle and the Nile Delta, Niger Delta, Lake Victoria, Lower Zambezi River (Thieme 7.7 Conservation recommendations 7.7.1 Planning and Policy Conservation of aquatic plants can be facilitated through and systematic planning processes policies effective for plant important areas value and protect that identify, conservation. Having identifi is for management actions important policy requirement at the catchment scale, employing such to be targeted 222 CHAPTER 7 | PLANTS species. based conservation atanyoftheselocationswill benefi in LesothoandKwaZulu-Natal province inSouthAfrica.Site Africawheresouthern itisrestricted toonlythree locations Rufi is to thewildfollowingsitebased restoration actions(see being recovered is species whichmightbenefi 2010) butseeChapter8,thisvolume.Anexampleofa sites havebeenidentifi et al. of speciesinimminentdangerglobalExtinction(Ricketts places thatharbouracriticalproportion ofthepopulations of sitebasedconservationactions.AZEsitesare specifi Extinction (AZE)provides agoodexamplefortherelevance X\]f\okèZk`fe ]ifd jXm\[ Aponogetonranunculifl Nympahea thermarum Nympahea thermarum ji FloodplaininTanzania. Jg\Z`\jèk_\jgfkc`^_kÆ 2005 ). Currently, 140speciespresent in82different ex situ ed inAfricaandMadagascar(AZE andcanhopefullybere-introduced , thisvolume) which isExtinctintheWildand orus t from sitebasedconservation © DAVID ALLEN (EN), which is endemic to (EN),whichisendemicto 8jdXccnfe[\if]eXkli\ . Another example Anotherexample t this c fi improving watershedsandwetlands, enhancingnatural water supplyand qualityandwastewatermanagement by an activepolicyoptionforaddressing challengesofhealth, recommendations is to “applyecosystemrestoration as restoring wetlandsfor thispurpose.Indeed,oneitsmain development, andspecifi Programme encourages restoration asatoolforsustainable a recent assessmentbytheUnitedNationsEnvironmental restoration ofdegradedwetlandecosystems.Forexample, wetlands issupportinganincreasing momentumforthe TEEB 2009).Thisincreased recognition of thevalue established (e.g.,MillenniumEcosystemAssessment2005 fl which, amongotherthings,servestoprevent oralleviate The importanceofvegetationasnaturalinfrastructure, 7.7.5 Restoration ltration” (Nellemann andCorcoran 2010). ood eventsandtopurifypollutedwaters,isnowwell cally pointsouttheimportance of CHAPTER 7 | PLANTS 223 ood cient species ed in Figure 7.7.ed in Figure eld survey and long-termeld survey ll these gaps. Data Defi ll these gaps. Data ve of all species assessed face risk of global Additional research is needed to increase our our is needed to increase Additional research knowledge for the 14% of species for which extinction risk assessments could not be completed due to lack of knowledge on their distributions, population status, and taxonomic status. One in fi extinction. One in four of all species endemic to continental with extinction. threatened Africa are coasts Central Africa, the Mediterranean and Atlantic and the Cape Fold Algeria and Tunisia, of Morocco, with the highest numbers of the regions are area, species. threatened and human expansion Habitat loss due to agriculture to is the main threat (settlements and infrastructure) aquatic plants. to aquatic plants pollution is a major threat Water mainly due to agricultural and domestic run-off. Development planning policy must include adequate consideration for the conservation of aquatic in development moves forward plants to ensure with minimal sustainable way, an environmentally possible. A impact to wetland ecosystems wherever combination of catchment and site scale actions is recommended. Aquatic plants play a major role in the provision the provision in a major role Aquatic plants play services including fl of many ecosystem waters and of polluted alleviation, bio-remediation to wetland also essential are nutrient cycling. They production of primary source ecosystems as the Their conservation is at the base of the food-web. importance. of great therefore The extinction risk of 718 aquatic macrophytes mainland continental Africa has been across assessed, including 365 species representing assessed families, and an 21 comprehensively another 44 families. additional 353 species from ➒ ➌ ➍ ➎ ➏ ➐ ➑ 7.8 Key messages ➊ ➋ to be completed (Beentje and Smith 2001; Ghogue 2010). 2001; Ghogue and Smith (Beentje completed to be status,taxonomic which the plants for many aquatic are There unknown. remain trends population size and distribution, fi taxonomic study, Additional will help to fi monitoring guided by as research, for such primary targets might be the identifi of DD species the concentrations ts ting oras are yetoras are 2008). ts for wildlife and local et al. nancially viable in many cases (TEEB nancially oras, and there are some countries for which fl are oras, and there recovered through restoration were valued at more than valued at more were restoration through recovered project. twice the cost of undertaking the restoration services that the wetland provisioning the Furthermore, estimated at EUR 315 per year in an area are now provides survive on an income of less 50% of households where than EUR 520 per year (Pollard Restoration is fi communities. For example, in South Africa a government- (WfW), carried out for Water Working funded programme, the Manalana wetland in Mpumalanga to restore a project in 2006. The site, which was degraded by erosion, important services to the local offer to was considered communities, with at least 70% of the people benefi was completed, an After the restoration in some way. economic study concluded that the livelihood benefi 7.7.6 Further research The availability of information on aquatic plants is relatively in some of the African with considerable gaps remaining low, fl 2009) and provides multiple benefi 2009) and provides Jg\Z`\jèk_\jgfkc`^_k 8jdXccnfe[\if]eXkli\ jXm\[]ifd\okèZk`fe Magdalena, C¹ and Juffe Bignoli, D²

ymphaea thermarum, Botanic Gardens, Kew, while not a

SPECIES IN THE SPOTLIGHT listed as Extinct in the single one has survived in the wild.

| Wild, is unique. It is the If Professor Fischer had missed smallest water lily in the the ”unusual” little plants while Eworld, endemic to one single site in he was looking at the area, the Rwanda, and has not been recorded species would never have been elsewhere, despite extensive discovered and its extinction would CHAPTER 7 searches. The intriguing story about have gone unnoticed. If Fischer this scientifi c rarity seems more had considered that his duty as a like a drama screenplay than a true botanist ended just as he described scientifi c achievement. the species instead of also trying It was discovered in 1987 by to fi gure out the conservation Professor Dr. Eberhard Fischer status, then the plants would have at Mashyzuza near Nyakabuye reached Germany only as dried in Rwanda. Its specifi c habitat (and therefore dead) specimens requirements seem improbable; it The minute fl ower of Nympheae and there would have been no is known only from a muddy site thermarum, listed as Extinct in the material from which to propagate. fl ooded by a series of hot springs Wild. © C MAGDALENA In addition, had the Bonn Botanic that fl ow at 80ºC, where the waters Gardens failed to provide facilities have cooled to approximately 24- and horticultural skills over a long 26ºC (Fischer 1993). the last 50 specimens growing in period of time, it would have also A few specimens were airlifted the wild died. Probably millions been lost forever. If Bonn had from its only known location of years of evolution in stable decided to keep these little gems for soon after its discovery and were conditions were wiped out in one themselves instead of sharing them kept at Bonn University Botanic single human action. with other institutions, then the Gardens, where botanists were In 2009, Carlos Magdalena, a mystery about how to propagate hoping to propagate the species. senior horticulturist from the Royal this species may have never been All attempts failed. While growing Botanic Gardens, Kew in London, solved at Kew. mature specimens seemed to be unveiled the secret. While the All the above highlights the relatively easy and seeds easily seeds of all other Nymphaea species importance of knowing what germinated, the seedlings were are traditionally grown deep, species we have, where they grow impossible to grow to maturity and submerged in water, this species and which ones are really in peril. died shortly after they germinated. was again different. Growing Once those are identifi ed, we can The urgency for understanding the seedlings in pots fi lled with spare the resources and the skilled its growing requirements became loam totally damp but so shallow staff able achieve the goal of increasingly evident as repeated that the surface of the compost keeping them going. This particular searches in numerous hot springs (and therefore the seedlings) was species has a huge potential for a across central Africa failed to fi nd exposed to the air, was the simple reintroduction programme, and another single population (Fischer, change that made all the difference its story is fascinating enough to pers. comm. 2010). In 2008, a pump and was all that was needed to save raise awareness about the wonders was installed in the only hot spring the species. Once its ex-situ growing and singularities of nature and, where N. thermarum used to grow, requirements were met and fully sometimes, its compelling fragility. sequestering the water before it understood the species was easy to N. thermarum illustrates a success reached the surface. The whole propagate. Hundreds of plants are story for which its last chapter is habitat dried out in the sun, and currently being grown at the Royal yet to be written.

¹ Royal Botanic Gardens, Kew. 224 ² Freshwater Biodiversity Unit, IUCN. CHAPTER 7 Jg\Z`\j`ek_\jgfkc`^_k |

JX`c`e^Õfn\ij SPECIES IN THE SPOTLIGHT Cook, C.D.K

agarosiphon is a genus with information that led to the species names. On top of it all, they are nine species that are confi ned being declared a noxious weed in not particularly good at producing to Africa and Madagascar. South Africa although it is in fact, oxygen. One species (Lagarosiphon endemic to southern Africa, confi ned When they fl ower a real wonder Cmajor) has become naturalized in to an area extending from southern takes place. The fl ower buds of the Europe and New Zealand. It has Zimbabwe to South Africa, where male plants become detached from been in Europe since, at least, 1910. it leads a modest and blameless the mother plant; they are liberated Occasionally it fl ares up in pools existence! under water and come to fl oat on the and lakes where it may interfere Although most species of surface of the water. Then, relatively with sport fi shing. In New Zealand Lagarosiphon are common, they have quickly and in front of your eyes, it has become a serious pest; large not merited presentable vernacular they open. At fi rst the sepals and populations can choke dams and names – they are just called oxygen petals bend back on themselves. rivers, interfering with the generation weeds or babergrass. Even the Latin Their outer surfaces are wettable, of electricity and irrigation. It has, name, which is in fact Greek, is so they stick to the water surface, therefore, been gazetted as a class boringly technical: Lagarosiphon is which stabilizes the fl ower so that it B noxious plant. Following this, it derived from the Greek words lagarós, cannot capsize. The inside parts of was declared to be a pest in Australia meaning thin or weak, and siphon, the fl ower are unwettable and remain (one can be fi ned up to AUS 80,000 tube - which refers to the long tube dry. Three fertile stamens then stretch for introducing it to Queensland) joining the ”fl ower” with the ovary. horizontally, parallel to the water and also in the USA, despite the fact From a distance the plants do not surface. Three sterile stamens become that it is not naturalized in either of look exciting, and perhaps deserve feather-like and elongate, eventually these countries. It is probably this their lack of interesting common reaching upwards to form a sail. The male fl ower is a perfectly engineered tiny sailing boat that skims over the water surface with the slightest breeze. This is something unique in the plant kingdom The stamen is also very special. Only one pollen mother cell from each of the four pollen sacs divides. Each stamen thus develops only 16 pollen grains. This is the minimum possible number of pollen grains for a complete anther. The anthers take a vertical position at right angles to the stamen fi lament. The pollen grains are abnormally large; they are presented on the outer surface of the anther, just in the right position to deposit their grains on the upright stigmas of the female fl owers, which they hit while skimming over the water surface. Just because Lagarosiphon major leads a modest life in southern Africa today, it does not mean that it must stay so. Perhaps major alterations to the environment could encourage its ability for very fast growth, and it Lagarosiphon major fl owers. © C.D.K. COOK could become a serious threat. 225 Jg\Z`\jèk_\jgfkc`^_k K_\_XY`kXkZi\Xkfi Sieben, E¹ almiet (Prionium serratum) choke the river leading to inundation is the only African of large areas of the valley. In this representative of the family situation, peat will start to form and Thurniaceae, but up to the the palmiet itself will then be able recent past it has been allocated to further expand the area that it SPECIES IN THE SPOTLIGHT G

| either to the family Juncaceae or occupies. In these situations, palmiet given its own family Prioniaceae. will create a habitat for many other Even though palmiet in itself is not riverine and wetland species that a threatened species (listed as Least grow between the palmiet plants. Concern), the unique ecosystems that The species occurs in two disjunct CHAPTER 7 it forms are becoming increasingly populations on two different rare. strata of quartizitic sandstone. The Palmiet is a special type of plant Table Mountain group sandstones in that it is an ecosystem engineer. It The palmiet (Prionium serratum) (LC) is are found in the Western Cape creates wetlands by growing in dense endemic to South Africa. It is common Mountains. Here Palmiet occurs stands that block the fl ow of rivers. in the Western Cape province in South together with plants such as These wetlands are subsequently Africa. © E. SIEBEN Wachendorfi a thyrsifl ora, Cliffortia colonized by many other plant strobilifera and Calopsis paniculata. species that would otherwise not On the Natal Group sandstones, fi nd suitable habitats. In that sense, protect the living tissues against fi re. extending from the Eastern Cape P. serratum forms the basal structure On top of the stem it carries a crown into southern KwaZulu-Natal, for a complete ecosystem, and is of stiff greyish serrated leaves with palmiet is also found, often occurring responsible for creating the most sharp edges, making it diffi cult to together with Pondoland endemic extensive peatlands of the Cape move around in an area dominated by species such as Leucadendron Floristic Region. Pools that come palmiet. Underground, the palmiet pondoense and Syzygium pondoense. into existence when river fl ows are plant has a dense network of fi brous The palmiet here has slightly blocked by Palmiet also provide roots that extend far deeper than narrower leaves with less of the valuable habitats for animals such as any other wetland plant occurring greyish shine as seen in the Table fi sh. For example, the Clanwilliam in its habitat, sometimes reaching Mountain populations, so it has been Yellowfi sh (Labeobarbus capensis) fi ve metres deep. This root network suggested that this may represent a is a threatened species (Vulnerable) extends deep into the peatland second species. The name ”Palmiet endemic to the Olifants River System and, since palmiet grows in dense, Valley” that can be found on maps on the West Coast of South Africa, widespread, stands, it is assumed of the city of Durban in southern where deep permanent pools with that this underlying fi brous root KwaZulu-Natal, suggests that the good cover provided by palmiet beds system has a considerable impact on distribution range of this second are important refuges for the species ecosystem functioning. form of palmiet historically must in seasonal rivers during the hot dry Palmiet occurs in two very distinct have extended further north than it summers. habitats: mountain streams and does today. Palmiet’s distribution is limited peatlands. In the fi rst habitat, small Most farmers in the area have to Table Mountain Sandstone plants of palmiet can be found a paradoxical relationship with (TMS). It only grows on this lining Cape mountain streams palmiet. They use water found in the extremely nutrient-poor substrate or even headwater seepages. The peatlands to irrigate their land, but so does not occur away from the second habitat is created entirely by simultaneously cut away the palmiet Cape Mountains. The nutrient- autogenous succession (where the plants themselves in effort to limit poor conditions in which palmiet stimulus for change is internal) of encroachment into their land. In the grows are in sharp contrast with the palmiet plants themselves. In the past, the leaves of the palmiet have the luxurious growth form that foothill zone of rivers the species can been used as fi bres, but this use is not it exhibits. A single ramet of the form dense stands that slow down very extensive nowadays. The fi bre, plant consists of a light woody stem the water fl ow. Since palmiet is called palmite, is the original source covered in old leaf sheaths, which able to grow in fl owing water it can of the name for this plant.

226 ¹ University of the Free State, South Africa CHAPTER 7 Jg\Z`\j`ek_\jgfkc`^_k NXk\i_pXZ`ek_#Xk_i\Xkkfk_\ |

]i\j_nXk\iY`f[`m\ij`kpf]n\jk\ie8]i`ZX SPECIES IN THE SPOTLIGHT Ouédraogo, R.L.¹ ichnornia crassipes, or water hyacinth, is an aquatic plant within the family Pontederiaceae and is

¹ Institute de l’Environment et de Recherches Agricoles, Burkina Faso 227 Chapter 8. Synthesis for all taxa

Holland, R.A.¹, Garcia, N.¹, Brooks, E.G.E.¹ and Juffe, D.¹, with contributions from: Howard, G.² (Section 8.2.3); Foster, M.N.³ (Section 8.3.4); Harrison, I.³, Upgren, A.³, Alonso, L.E.³ (Section 8.5); Matthews, J.H.4, Wickel, A.J.5, Freeman, S.5 and Thieme, M.L.5 (Section 8.6)

1 Freshwater Biodiversity Unit, IUCN Species Programme, Cambridge, CB3 0DL, UK 2 Global Invasive Species Coordinator, IUCN, Wasaa Conservation Centre, Mukoma Road, P.O. Box 68200, 200 Nairobi, Kenya 3 Conservation International, 2011 Crystal Drive, Suite 500, Arlington, Virginia 22202, USA 4 Current address: Conservation International, 2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA 5 WWF Conservation Science Program, World Wildlife Fund, 1250 Twenty-Fourth Street, N.W., Washington, DC 20037, USA

MAIN: Fishers return to Senanga on the Zambezi River, at the southern end of the Barotse fl oodplain. © JENS KIPPING RIGHT: High species diversity in a fi sh catch at Ilagala market, Lake Tanganyika, Tanzania. © KLAAS-DOUWE DIJKSTRA CONTENTS 8.1 Introduction 230 8.2 Freshwater biodiversity across Africa 230 8.2.1 Patterns of richness and main threats 230 8.2.2 Distribution of threats to African freshwater species 235 8.2.3 Biological invasions affecting freshwater species in Africa 244 8.3 Conservation action 247 8.3.1 Congruence between taxonomic groups 247 8.3.2 Protected areas for freshwater ecosystems 247 8.3.3 Key Biodiversity Areas for freshwater species 249 8.3.4 Alliance for Zero Extinction – AZE sites 249 8.4 Livelihoods dependence on freshwater species 251 8.5 Areas for further work 258 8.5.1 Regions of Data Defi cient species and threatened species 258 8.5.2 Comparing distributions of Data Defi cient species with records of new species. 262 8.6 The future of African freshwaters 264 8.6.1 Climate change and African freshwater conservation: seeing the future through the past 264 8.6.2 African waters: special qualities and challenges 265 8.6.3 A short history of African eco-hydrological dynamism: the past 100,000 years 265 8.6.4 Signs of anthropogenic climate change in Africa 267 8.6.5 Projecting trends and responses 267 8.6.6 Implications for facilitating healthy responses to freshwater change 268 230 CHAPTER 8 | SYNTHESIS FOR ALL TAXA face. Africa ishometo asignifi and considerone ofthemostsignifi effort. Finally, weexaminethefuture ofAfricanfreshwaters as theserepresent importantfocalareas forconservation Biodiversity Areas and AllianceforZero Extinctionsites, freshwater biodiversity andidentifythelocationofKey the role thatexisting protected areas haveinconserving Weand considerwhatdrivesthese patterns. discuss of richnessandthreatpatterns across Africancountries all fi continental Africa.We present acombinedanalysisfor distribution offreshwater biodiversityacross mainland presented inchapters3to7andconsiderthestatus In thissynthesischapter, wecombineinformation 8.1 Introduction Platythelphusa maculate sh, molluscs,crabs, odonatesandplants,toillustrate (LC),asmallcrabendemictoLakeTanganyika, hidesinempty cant proportion ofthe cant threats thatthey continued survivalandprovision ofbenefi and theactionsthatneedtobetakeniskeyensuringits and statusofthisbiodiversity, thethreats whichit faces (Table 8.1)and,assuch,understandingthedistribution world’s speciesthatare dependentonfreshwater habitats relevant forpolicy makers. atthecountrylevel tohighlightimportantpatterns patterns unit forfreshwater species.Here, wefi sub-catchment scale,asthisrepresent thelogicalmapping In previous chapters, analysishasbeenconductedatthe ofrichnessandmainthreats8.2.1 Patterns 8.2 Freshwater biodiversityacross Africa Neothauma shells. © SASKIAMARIJNISSEN ts topeople. rty consider rstly CHAPTER 8 | SYNTHESIS FOR ALL TAXA 231 cant numbers rst described it for ed as being of conservation concerned as being of conservation At the country scale, richness for all freshwater groups groups At the country scale, richness for all freshwater Such a pattern the equator. from decreases 8.1 in Table a general rule in ecology has been suggested to represent H. Rapoport, who fi named after Eduardo Rule has been called into mammals. Although Rapoport’s years (Gaston 1998), patterns question in recent shown in that, at this course scale, richness 8.1 demonstrate Figure is highest in equatorial countries and taxa of freshwater with latitude. decreases Equatorial countries also contain signifi of species classifi 8.2), with the highest Red List (Figure to the IUCN according of the centre concentrations occurring in a band across D. R. Congo, Uganda and Tanzania, the continent through species are high numbers of threatened Kenya. However, 19% 12% c. c. % of global total found in Africa Number of species assessed in Africa Number of described species MolluscsOdonatesAmphibians >5000 5680Waterbirds 4251Mammals 868 624 705 145 624 279 12% 32 15% 32% 22% CrabsFish 1280 >15000 120 2836 9% Taxon Taxon Group Table 8.1. Estimated number of inland water 8.1. Table dependent species by major taxonomic group. Species - Red List of Threatened the IUCN Data from September 2010. Figure 8.1. Richness of species across continental Africa based on occurrence in countries. in countries. occurrence based on Africa continental across species of 8.1. Richness Figure 232 CHAPTER 8 | SYNTHESIS FOR ALL TAXA Figure 8.2.Richnessofthreatened speciesacross continentalAfricabasedonoccurrence incountries. species in theGreat Africa, riversincoastal Lakesofeastern areas of highestrichnessare drivenbythediversityof between countries emerges. Across equatorial countries, picture withinand ofthedrivers ofbiodiversitypatterns molluscs andplantsatthesub-catchment scale,aclearer By combininginformationon fi high numbersofthreatened speciesinthiscountry. resources andthespread ofinvasivespecieshasledto south oftheequator, developmentofSouthAfrica’s water production inthiscomparativelyaridlandscape.Moving over-abstraction ofwaterassociatedwithagricultural has highnumbersofthreatened species,mainlydueto Nigeria). To thefarnorthofcontinent,Morocco also (Mozambique, Malawi)andupthewestcoast(Cameroon, also foundincountriesextendingdowntheeastcoast sh, odonates,crabs, habitats. When these temporary water bodies dry up, the habitats. Whenthesetemporary water bodies dryup, the that exploittemporary poolsorevenpuddles in roads as known are theannualKillifi itself inmudwhenitsfl ( the mostfamoussuchadaption isfoundintheLungfi arise through fl that enablethemtocopewithever changinghabitatsthat Africa ishometosomespecieswithremarkable adaptations few suitablehabitatsforfreshwater species. the Sahara,Namib,andKalaharideserts,where there are of lowestrichnesscorrespond witharidregions, suchas the continent,primarilyinD.R.Congo(Figure 8.3).Areas that tracethecourseofmajorriversthrough thecentre of Africa,andsub-catchments sub-catchments inwestern Protopterus annectens ooding or seasonal rainfall patterns. Perhaps ooding orseasonalrainfallpatterns. oodplain habitat driesup.Lesswell (LC)), which is able to cocoon (LC)),whichisabletococoon s (e.g., sh Nothobranchius spp.) spp.) sh sh CHAPTER 8 | SYNTHESIS FOR ALL TAXA 233 ) sh Protopterus annectens Protopterus ood plains, and when The African lungfi ( (LC) normally lives on fl these dry up it survives by secreting a thin slime around itself, which dries into a protective cocoon. © ROGER BILLS/SAIAB Species richness = number = number richness Species ed, broadly ve freshwater groups at the sub-catchment scale. the sub-catchment at groups ve freshwater ndings from other studies ndings from rmed fi sh can remain buried in the mud for years until buried sh can remain water once again returns to awaken them. species freshwater where In the most arid regions, is often intense coexist with human populations, there of high areas creating on water resources, pressure species. For example, in concentrations of threatened sub-catchments containing high and Algeria Morocco situated along the species are numbers of threatened the majority of the human population coastline where the continent a similar pattern is found. Across is often of to areas richness corresponding found with threat 8.4). Although a high human population density (Figure identifi are drivers of threat range of different confi speaking, results that activities associated with agricultural production eggs of the fi Figure 8.3. Combined richness of fi richness 8.3. Combined Figure sub-catchment. river/lake per of species 234 CHAPTER 8 | SYNTHESIS FOR ALL TAXA scale. Figure 8.4.Combinedrichnessofthreatened speciesbasedonthefi river networks, often crossing international boundaries and river networks, oftencrossing boundariesand international have thepotential topropagate widely throughout lakesor where activitiesinonepartofalake orrivercatchment issues surrounding the managementoffreshwater systems, species intheselakeshighlight oneofthemostdiffi lakes, threatening many species.Theproblems facedby have signifi such asoverfi and Malawi.Asdiscussedinprevious chapters,pressures, arises asthecountryborders LakesVictoria, Tanganyika of threatened speciesrecorded forTanzania (Figure 8.2) Based onsub-catchmentanalysis,thehighconcentration freshwaters (Allan2004). pervasive threat tospeciesandthehealthofworld’s and urbanisationwithincatchmentsrepresent themost Threatened richness=numberofthreatenedspeciesperriver/lakesub-catchment. cantly impacted the rich endemic fauna of these cantly impactedtherichendemicfaunaofthese shing, invasivespeciesandeutrophication, cult cult to protect aquatic biodiversity. example where countriesmustwork withtheirneighbours case oftheGreat Lakeshighlightedhere represents justone not beaddressed by countriesactingontheirown.The to address problems relating towaterresources thatcould operation Potential’fostersco-operation betweencountries the UNESCOinitiative‘From PotentialConfl been demonstratedinanumber ofinstances,forexample, fostering co-operationtoprotect shared resources. Thishas involved inmanagementactionshastheaddedbenefi freshwater systemsmeansallstakeholdersneedtobejointly of waterresources. Realisationthattheconnected nature of due tosuchtransboundaryissues,principallyrelating touse countries. However, there isalonghistoryofdisputesarising requires aco-ordinated approach tomanagementbetween management zones.Thetransmissionofthreats inthisway ve freshwater groups atthesub-catchment ict toCo- t of of t CHAPTER 8 | SYNTHESIS FOR ALL TAXA 235 ve ve % Threatened © SASKIA MARIJNISSEN ed for African freshwater species, these species, these ed for African freshwater NT LC DD Documenting threats is important for the Red List, not is important for the Red List, not Documenting threats also for guiding just for assessing species’ status, but chapters conservation action. Previous appropriate to the fi a detailed discussion of threats provide taxonomic groups assessed. Currently, spatially explicit spatially explicit assessed. Currently, taxonomic groups is not collected as part of data on the location of threats the distribution the Red List assessment. By examining and identifying where threatened, of species that are of the total high numbers of species or a high proportion we attempt to identify priority threatened, species are and conservation action. For for further research areas to the main each sub-catchment, analysis was limited identifi threats logging, water agriculture, include water pollution through harvesting for extraction, habitat loss due to agriculture, development, food at subsistence level, infrastructure mining, sedimentation, and invasive alien species. The varies between taxa, so in some importance of threats instances we describe patterns based on a subset of of particular concern, to highlight areas species in order instances our analysis is based on in other whereas combined data for all groups. global scale by taxonomic groupings. cations at the 9% 12% 2% 55% 20% 23% 5% 13% 3% 57% 19% 22% 9% 12% 2% 55% 20% 9% 5% 13% 8% 49% 18% 25% EN VU 12% 7% 6% 34% 30% 29% 2% 4% 9% 2% 6% culty. With the development culty. ed as threatened, according to IUCN according ed as threatened, ects this diffi ve newly assessed freshwater groups, 21% of groups, assessed freshwater ve newly Distribution of threats to African freshwater to African freshwater Distribution of threats species Taxonomic Taxonomic group EX EW CR Crabs 0 0 Fishes 0.1% 0% Molluscs 3% 0% Odonates 0 0 Plants 0 0.2% Table 8.2. Summary of the IUCN Red List Category classifi 8.2. Table Red List Categories and Criteria. The level of threat is Red List Categories and Criteria. The level of threat and lowest highest for mollusc species (29% threatened) 8.2). (Table for odonates (9% threatened) species were classifi species were may lead to changes in the status the IUCN Red List September 2010. Subsequent revisions from Data current by changes in these statistics. of species which will be refelcted of rich mineral deposits to the east of the region, and the region, of rich mineral deposits to the east of the species, the establishment to freshwater associated threat status, against which to of a baseline for species current priority. impact, should be an urgent measure 8.2.2 the fi Across While the high number of threatened species in the Great species in the Great While the high number of threatened for emerges Lakes is of concern, positive picture a more the the rich fauna of rivers through central Africa, where species few globally threatened Congo contain relatively As highlighted in to their overall biodiversity. in relation a quite different presented chapters, this area previous due to the challenge during the course of the assessment, lack of information. The high number of DD species relative refl in this region Erosion and sedimentation, as seen here, pose a major threat freshwater to many species. 236 CHAPTER 8 | SYNTHESIS FOR ALL TAXA sub-catchment. species perriver/lake number ofthreatened Threatened richness= (c) sedimentation. invasive species;and to: (a)agriculture; (b) as threatened due of fi Figure 8.5.Thenumber sh speciesclassifi ed (b) (a) CHAPTER 8 | SYNTHESIS FOR ALL TAXA 237 holds

known known

sh cant threats . 2010). Our analysis Haplochromis crebidens, Haplochromis crebidens, et al . sh from Lake Victoria and and Lake Victoria sh from species: H. vittatus cant pressures on species in the Great on species in the Great cant pressures and Haplochromis threatened by harvesting for subsistence, including for subsistence, including by harvesting threatened

sheries across Africa (Dugan sheries across H. occultidens, H. olivaceus, H. paucidens, H. rubescens, H. occultidens, H. olivaceus, H. paucidens, H. rubescens, H. scheffersi one of the most signifi represents Agriculture an interesting This represents to aquatic systems globally. of catchments, as increasing for management trade-off by the application of fertilisers land based food production and pesticides can have a negative impact on the harvest within the catchment. freshwaters of food species from particularly threatened In the Lower Guinea, species are by the negative impact of activities such as intensive banana, and palm oil rubber, land use and pollution from to be impacted recorded plantations. Up to 29 species are the and all (21 threatened), agriculture by habitat loss from by water pollution arising affected are species threatened activities. The eastern agricultural from slopes of Mount and Middle Mungo River) is an area (Upper Cameroon range and restricted of threatened high numbers where of many people in sub-Saharan Africa. There is growing is growing of many people in sub-Saharan Africa. There of the largest evidence of a decline in the health of some fi indicates signifi with many fi Lakes region, Lake Kivu by over-harvesting. Malawi threatened fi range freshwater a high richness of restricted seven to be sh ed into sh have diversifi sheries for the people of sh diversity) are under risk under risk sh diversity) are sh form a vital component of the diet sh form a vital component of the diet sh fauna being of particular concern. Lake (c) diversity across the continent, are threatened due to to due threatened the continent, are diversity across 8.5a), 23 species (Figure agriculture water pollution from total fi (4% of the continent’s of extinction due to invasive species (Figure 8.5b; but see 8.5b; but see of extinction due to invasive species (Figure a scapegoat?), and 29 Chapter 1, Box 1.1 Is Nile perch due threatened total) are species (5% of the continent’s 8.5c). The changes seen in Lake to sedimentation (Figure of particular concern, past decades are over the Victoria due to the importance of inland fi fi Africa. Freshwater Results of the analysis show that major concentrations Results of the analysis show that major species occur along the freshwater of threatened Algeria Mediterranean and Atlantic coasts of Morocco, southern the Upper and Lower Guinea, and and Tunisia, in easterneastern Lakes the Great South Africa and priority regions represent Africa. As such, these areas stands out as hosting for conservation. Lake Victoria species with number of threatened high a remarkably its unique fi as it dreampond’ dubbed ‘Darwin’s has been Victoria most spectacular demonstration the world’s represents of evolution, as just a handful of fi of species, possibly over a period of many hundreds just 15,000 years, meaning that a new species arises on average every 30 years (Spinney 2010). 2% of the total fi Thirteen species, representing 238 CHAPTER 8 | SYNTHESIS FOR ALL TAXA and Lambin 2002).The lossoftree coverand associated institutional, technological anddemographicfactors (Geist highest intheworld, drivenbytheinterplayof economic, Rates ofdeforestation across Africaare amongstthe freshwater andterrestrial, arisesthrough deforestation. One ofthemostcomplex threats tospecies,both vineyard development. agricultural activities,includingcitrus,deciduousfruitand richness =numberofthreatenedspeciesperriver/lakesub-catchment. Figure 8.6.Thetotalnumberofspeciesforthefi restricted range Cape region alsoholdsparticularlyhighnumbersof due toagriculture. TheOlifantsRiverbasininWestern and threatened freshwater fi species canbefound,includingupto21restricted range

fi sh andodonatathreatened byintense sh impactedbyhabitatloss ve groups classifi Upper Guinea andMount Cameroon. identifi 1997). Deforestation associatedwith agriculture was where forests are disappearingatthefastestrate(FAO This isconsidered to betheregion intropical Africa of the of agriculture expansion, iswidespread alongthebanks and Tanzania. Africa,deforestation, Inwestern asaresult coastofMozambique Africa andalongtheeastern western deforestation threatens highnumbersofspecies incoastal thermal regimes (Allan2004).AsillustratedinFigure 8.6, changes tobiogeochemicalprocesses, hydrology and on thefunctioningoffreshwater systems,including disturbance ofthecatchmentcanhaveprofound affects

ed asaprincipal threat tocrabsandodonates in ed asthreatened bydeforestation. Volta River

at AkosomboinGhanaand Niger. Threatened CHAPTER 8 | SYNTHESIS FOR ALL TAXA 239 Threatened Threatened cant impact on downstream freshwater © JANE BOLES Rates of deforestation across Africa are amongst the highest in the world. Loss of tree cover from within a catchment can have a signiﬁ ecosystems. abstraction. due to water ed as threatened

where many where

ict. Within such . 2010). As illustrated using freshwater . 2010). As illustrated using freshwater et al wells, which are the sole known habitat of some species, the wells, which are out. Perhaps the most vivid example at risk of drying are Lake arises from placed on water resources of pressure Chad, which has shrunk by an estimated 95% since molluscs (Figure 8.7), water abstraction represents a 8.7), water abstraction represents molluscs (Figure Atlantic Plains of Morocco, in the major threat Threats to biodiversity attributable to competition for to biodiversity Threats the continent vary considerably across water resources (Vörösmarty disagreements, the allocation of water for maintenance of the disagreements, (Dudgeon 2006). considered aquatic biodiversity is rarely Competition for water resources has become an Competition for water resources has, decades and issue in recent prominent increasingly in armed confl in some instances, resulted richness = number of threatened species per river/lake sub-catchment. river/lake per species of threatened = number richness Figure 8.7. The total number of mollusc species classifi mollusc species of total number 8.7. The Figure 240 CHAPTER 8 | SYNTHESIS FOR ALL TAXA remarkably high numbersoffreshwater speciesthreatened mineral resources canhave onspeciesisrefl value ofthedeposits. Theimpactthatexploitation of has beenintenselymodifi highlands oftheNimbaregion, where thelandscape of species.Molluscsare particularlyimpactedinthe (bauxite), goldanddiamondspose athreat toanumber Cameroon, open-pit mining activitiesforoil,aluminium species withintheGuineaBasin deposits hasresulted inaconcentrationofthreatened Africa,exploitationofareasIn western richinmineral (Chapter1). changing rainfallpatterns the mid1960sduetowaterextractionforirrigationand richness =numberofthreatenedspeciesperriver/lakesub-catchment. Figure 8.8.Thetotalnumberofspeciesforthefi ed asaresult ofthe economic . In Nigeria,Ghanaand ected inthe ve groups classifi systems (Allan2004). The rapidgrowth of large cities, can increase thelevelofpollutantsthat reach freshwater profoundly infl river systems(Richter species andprofoundly alterthecharacteristicsof dams, whichactasbarriersto migrationforfreshwater river systemshavebeenheavilyfragmented byover1million is amajorthreat toAfrican species(Figure 8.9).Globally, Construction ofdamsandotherinfrastructure development range species. in agriculture, isamajorcauseofthreat for many restricted Delta, togetherwithwaterpollutionfrom pesticidesused in theNigerDelta(Figure 8.8).OilexplorationintheNiger ed asthreatened duetomining. uences thehydrology ofcatchmentsand et al . 2010).Similarly, urbanisation Threatened CHAPTER 8 | SYNTHESIS FOR ALL TAXA 241 ji and Kilombero Rivers ji and Kilombero 2007). et al. cant impact on function within the ecosystem ed as threatened due to infrastructure due to infrastructure ed as threatened cant in the Cunene basin in Angola, the Atlantic plains on this approach, the Maghreb region in northern region Africa the Maghreb on this approach, of species with the highest proportion as the area emerges of intensive agriculture This is a result threatened. currently development and water extraction infrastructure activity, the impacting a comparatively limited fauna. Similarly, impact of invasive alien species is seen to be particularly signifi of northern the Lower Rufi Morocco, of a high proportion and the Niger in Mali, where in Tanzania where In those areas the overall biodiversity is threatened. can the loss of a single species overall biodiversity is low, have a signifi by the and so impact the goods and services provided (McIntyre freshwaters regions classifi ve groups . 2011). et al sh species through sh species through c threat, analysis of the proportion analysis of the proportion c threat, Threatened richness = number of threatened species per river/lake sub-catchment. per river/lake species threatened of = number richness Threatened In addition to consideration of the total number of species impacted by a specifi can threatened of species within sub-catchments that are for a high priority that are into areas useful insights provide 8.10. Based conservation action, as illustrated in Figure such as Freetown and Monrovia, in the coastal zones of and Monrovia, such as Freetown western urbanisation along the Niger Africa, and increasing Delta and the coastal plains of Sierra Leone and western by a hotspot of species threatened Liberia, has created urbanisation at Similarly, development. infrastructure of Kinshasa in D. R. Congo, the Pool Malebo, upstream fi of freshwater a number threatens and lead toxicity (Brooks sewage discharge development. development. Figure 8.9. The total number of species for the fi species for of total number 8.9. The Figure 242 CHAPTER 8 | SYNTHESIS FOR ALL TAXA and (d)invasivespecies. loss duetoagriculture; abstraction; (c)habitat water pollution;(b) groups threatened by:(a) from alltaxonomic proportion ofspecies Figure 8.10.The (a) (b) CHAPTER 8 | SYNTHESIS FOR ALL TAXA 243 (d) (c) 244 CHAPTER 8 | SYNTHESIS FOR ALL TAXA © GEOFFREYHOWARD species makestheircontrolespeciallydiffi range throughoutAfrica.Thehighcommercialvalueofsuch which hasbeenwidelyintroducedtosystemsoutsideitsnative Roadside saleof are notnative tomanyparts ofthecountry in whichthey system oradistinct habitatwithinalake,and assuch, African freshwater speciesare restricted toasingle river meaning notnativetoa rather thantheusualunderstanding of‘alien’or‘exotic’, this defi considering freshwater biodiversityinAfricatoensure that species anditsfunctions.Itisparticularly importantwhen causing damageofsomekindtotherecipient habitat,its refer tospeciesasan‘invasive’unlessthatinteractionis –soitdoesnotmakesenseto and thehabitatconcerned the result ofaninteractionbetweentheincomingspecies (CBD DecisionVI/232002).Biologicalinvasionisalways established, spread andcausedamagetobiodiversity being introduced toanewhabitat,where theybecome Biological invasionsare causedbyalien(orexotic)species what wemeanbyaninvasivespecies. examining thesecasesindetailitisimportanttodefi notorious examplesfrom across Africa.However, before on freshwater systems,weconsidersomeofthemost To illustratethedamagethatsuchspeciescanhave the oneofmaincausesbiodiversitylossglobally. biodiversity. Invasivealienspeciesare considered tobe species emerged asoneofthekeythreats tofreshwater During theAfricanfreshwater assessments,invasive 8.2.3 in Africa Biological invasionsaffecting freshwater species nition refers topresence orabsence ina Oreochromis country species, possibly . Thisisbecausemany cult. cult. O. niloticus habitat , ne

see in itsmanyformsasahighlyproductive aquaculture species: manipulated typesandstrains),whichisstillbeingpromoted at leasttworecognised sub-speciesandmanygenetically freshwaters istheNiletilapia, One ofthemostnotoriousexamplesfoundinAfrican for theentryofspeciesintonewhabitats. escapes from fi almost everyriverandlakebasinacross thecontinent, More recently, astheaquaculture tradehasexpandedinto yields ortoprovide enhancedsport-fi rivers, lakesandwetlandsinorder toenhancelocalfi non-African species,havebeenintentionallyintroduced to Freshwater fi invasive specieshaveentered freshwater habitats. Across Africathere are avarietyofroutes through which the habitatandimpactbiodiversity. species thencolonisesnewareas, where itmaydamage or through thetranslocationofspeciestonewareas; the country are removed, forexample,bylinkingwaterbodies their countryoforiginifbarrierstomovementwithinthe are found.Speciescantherefore becomeinvasivewithin (indigenous) of freshwater invertebrates.Ithashybridisedwithother essential aquaticplants,otherspeciesoffi is capableofcausingseriousdamagetopopulations 8]i`ZXÆX]i`\e[Xe[]f\ and will hunt larvalfi freshwater species.Bothspeciesare capableofomnivory areas, puttingpressure onmanyendemicandthreatened basins), andindicationsare thattheywillmovetoother Edward, theSemliki River, andtheNileZambezi lakes andriversofAfrica(forexample, LakesVictoria and These twospecieshaveinvaded someofthemajor locally andescapedinfested riverandlakebasins. Africa,whereand eastern theyhavecauseddamage to otheraquaculture farms,damsandlakesinsouthern Louisiana crayfi claw crayfi ago byfi (intentionally andlegally)tothecontinentseveraldecades alien freshwater crayfi serious problem inthefreshwater habitats ofAfricais Another categoryoffooditemthathasbecomea and, assuch,hassomecommercial value. and isthebasisofanexportindustryinsomecountries item inmanypartsofAfrica(fresh, frozen, smokedordried) and aquaculture species.Niletilapiaisthepreferred food its spread inthewild,asitissuchahighly-desired fi recorded inmanycases),itisespeciallydiffi invasive (seriousimpactstonativebiodiversityhavebeen is argued bysomethatthisspeciesdoesnotalwaysbecome even more damagingthantheirinvasiveparent. Althoughit different formsandresulting inhybrids thatare reportedly

sh farmers,andtwoofthose(the Australianred gZ\`k\ gk`_Æ jgfkc`^_k k_\ è Jg\Z`\j sh, sh nativetoAfrica,togetherwithseveral Oreochromis Oreochromis sh farmshavebeguntorepresent aroute Cherax quadricarinatus sh, Procambarus clarkii sh andfi sh. Fourspecieswere introduced . Asanomnivore, theNiletilapia species, furtherexpandingits ngerlings, other crustaceans, Oreochromis niloticus Oreochromis shing opportunities. shing `X``\Xjk\ie è K`cXg`X andtheAmerican ) were thenmoved sh andarange cult tomanage shery shery (with (with

CHAPTER 8 | SYNTHESIS FOR ALL TAXA 245 . liculoides Eichhornia Azolla fi oating alien aquatic Salvinia molesta, Pistia © GEOFFREY HOWARD oating mats, sometimes oating islands’ (different oating islands’ (different sh and water plants. The presence The presence sh and water plants. oating species), and has since become ora and fauna through direct as well as indirect as well as indirect direct ora and fauna through oating-leaved species and riparian or semi- ) has been able to occupy an apparently vacant (which has been in some parts of Africa for much other aquaticoating complex of sedges, grasses and cantly (Howard and Harley 1998). This, in turn, affects cantly (Howard oating mats which can support riparian vegetation, suchoating mats which can support riparian plankton to the largest fi plankton to the largest alter freshwater of water hyacinth has the potential to plants changes habitats such that the balance of component signifi now dominated by which are organisms, other freshwater conditions. new complexes of vegetation and environmental hyacinth to formA classic example is the ability of water fl that a fl and semi-aquatic species forms ‘fl the classic ‘sudd’ formations) which transport species from new invertebrates, create to new localities, contain different and, for a range of aquatic organisms and niches refuges disadvantage some endemic and threatened inevitably, hyacinth was joined during the early to species. Water other free-fl mid 20th century by three – tropics the American plants from stratiotes but is not especially invasive) and longer, Introduced in the 19th century, water hyacinth ( Introduced in the 19th century, crassipes niche (as a free-fl invasive throughout much of Africa. of forming fl also capable These are sometimes in combinations of the three separately, impacts as species, with similar (but not quite so severe) many other examples of invasive are water hyacinth. There species, emergent aquatic plants, including submerged species, fl competitors aquatic species. These plants can be effective (for light, nutrients, space, and so on) in particular habitats native species of both and can affect of Africa, and regions fl freshwater impacts, such as changed habitats and opportunities for and changes in water physical diseases, new herbivores, and chemical parameters. et (Hill

) sh sh, including sh, oating aquatic sh, meaning that ) and common carp ) and common Eichhornia crassipes sh species are capable of capable are sh species ted from a lack of competition ted from has altered the vegetation and and the vegetation has altered

sh ora and fauna of areas that they ) (pictured here) invasive have become ). Both crayfi Micropterus salmoides Micropterus sh, such as the Louisiana crayfi © G. HOWARD cantly alter the fl 2008) – assisted by a number of exotic fi by a number of 2008) – assisted owers, and because it gave a green covering to water covering to green owers, and because it gave a . 1998) that has been introduced into a continent that does into a continent introduced . 1998) that has been Cyprinus carpio Procambarus clarkia Procambarus al free-fl of large, not have any other species to Africa several times plants. The species was introduced water plant, forduring the early 19th century as an attractive its fl settlers liked to see and imagined bodies that early foreign evaporation/water loss. This extremely was reducing invasive species has benefi pests and diseases, and has and absence of controlling almost every now infested most rivers and wetlands and hyacinth covers Africa. Water water body in tropical large light and oxygen the water surface, leading to diminution of biodiversity from beneath, and consequently impacts local molluscs and insects, and devour aquatic vegetation. vegetation. aquatic devour and and insects, molluscs crayfi The Louisiana invertebrate fauna of Lake Naivasha in Kenya (Clearwater in Kenya (Clearwater fauna of Lake Naivasha invertebrate et al. ( bigmouth bass ( be so their impacts may watersheds, travelling between and invasion unless their spread in the future much greater for the success of these curtailed. The principal reason are of endemic crayfi two species is the lack and own group, within their from they face no competition native crabs. from only limited competition similarly contributed to the success Lack of competition has water hyacinth ( of the tropical in many freshwater systems across Africa; in part due to lack of competition. These species are omnivorous and signifi Freshwater crayfi colonise. ( 246 CHAPTER 8 | SYNTHESIS FOR ALL TAXA Even for well-studiedinvasive species, itisdiffi of namedexamples withintheproject’s regional reports. available (across mosttaxa),although there are a number may beaffecting threatened andendemicspeciesisrarely detailed informationontheprecise introduced speciesthat and are probably havingimpactsonnativespecies.But gastropods) havebeen recorded inAfricanfreshwaters Several speciesofalienmolluscs (bothbivalvesand quite tolerantofextendedperiodsinundation. riparian reeds andsedgesbygrowing over them,andis and plantsinAfrica.Itisalready threatening swampand an increasing threat tofreshwater speciesofanimals problem inpartsofAustraliaandAsiacouldbecome habitats. ThisNewWorld speciesisalready aserious space onfl countries, thetwomimosaspeciesare competingfor areas andsimilarlybecominginvasive–inatleasttwo invisa Mimosa diplotricha closely related (anddefi an almostcompletedearthofotherspeciesplants.A fl on arangeofspecies,from semi-aquaticantelopesto semi-aquatic habitathasalready haddeleteriousimpacts alien species).Suchadrasticchangeofaquaticand herbivores (areason forassumingthatitisindeedan vertebrates, whilesupportingveryfewinvertebratesas thickets andprevents entryofmostterrestrial andaquatic thousand hectares. Itexcludesallotherplantsfrom these shrubs (upto5mhigh)sometimescoveringseveral species toadense,monoculture oftightly-packedtall can changefrom beingasparselydistributedriparian wetlands across thecontinent(Shanungu2009). in smallandlarge fl This specieshasbecomeinvasiveinthelastthree decades World buthasbeenrecorded inAfricaforseveralcenturies. spiny shrubthatprobably originatedinthetropical New sometimes calledthegiantsensitiveplant,isaspeciesof in manyareas thataffect freshwater biota. leguminous shrubinparticularthathasbecomeinvasive Africa there isonespeciesofriparianandfl compressed intoanarrow area closetothewater. Across zone, whichoftenhasacomplexfl or evenextirpation,andthisisespeciallysointheriparian habitats ofmanyspeciescanresult inpopulationdeclines conservation habitatsandecosystems.Subtlechangesin and othervegetationtypesinbothproduction and widespread inAfricaandaggressively invadewoodlands Australian blackwattleandotheracaciasare now sites forseveralOdonataare nolongeravailable. The several streamsAfricasothattheperching insouthern ( freshwater species.TheinvasiveAustralianblack wattle both semi-aquaticandterrestrial plantsaffecting native riverbank habitatsare alsosusceptibletoinvasion by Occasionally inundatedfl odli fi oodplain Acacia mearnsii ), isalsospreading insomeriparianandfl sh andwaterbirds, and isassociatedwith oodplains andotheroccasionally-inundated ) hasaltered theriparianvegetationof (sometimeserroneously called oodplains andseasonally-inundated nitely alien)speciesofspinyshrub, oodplains andlake-edge ora andfaunathatis Mimosa pigra oodplain oodplain M. pigra Mimosa cl to cult ,

animals. whichsupportfewotherspeciesofplantsor monocultures dense many wetlandssystemsinAfrica,oftenforming Mimosa specieshavebecomeinvasivethroughout other species, asinvasive speciesimpact theirhabitat. may leadtoadeterioration oftheconservation statusof in increased pressure onalready threatened speciesand and otherchecksbalances. Failure todosomayresult freshwater habitatswithout therequisite riskassessments spread, andtoactively prevent furtherintroductions to affecting Africanbiodiversity, toreduce thechancesof therefore, to understandtheinvasionsthatare presently Patronshi 2005)–butthiswilltaketime.Itisimportant, hormones forsomeaquaticanimals(Kapuscinskiand genetic manipulationanddevelopmentalreproductive control. Newideasare slowlyemerging forcontrol using will notbespecifi yet more organisms, asthere isoftenafearthatthey there isoftenoffi proven tobeeffective (WilsonandGarcia 1992).However, invasive originates),andanumberhavebeenused using agents(specifi alien plants,there are usuallypossibilitiesforbiocontrol value andsoreduces thechancesofeffective control. For process ofusingsuchspeciesgivesthemaneconomic species through utilizationistempting,butthesimple means are oftenunsatisfactory. Managementofinvasive to usechemicalmethodsofcontrol, andmechanical 2003). Clearly, inaquaticsituationsitisextremely unwise – isextremely diffi alien species–especiallythosethatare mainlysubmerged the world.Findingsolutionstofreshwater invasionsby that isslowlydevelopinginanumberofcentres around This isthepurposeof‘InvasionBiology’,ayoungscience invasions, andsoreduce thethreat tonativeAfricanspecies. information topredict andsoprevent ormanagefuture the connectionsmore precise inorder togatherenough threatened speciesareitisessentialtomake concerned, pinpoint theexactnature ofnegativeimpacts.Butwhere © GEOFFREYHOWARD c tothespeciestheyare designedto cial resistance totheintroduction of cult atpresent (Howard andMatindi c pestsanddiseasesfrom where the CHAPTER 8 | SYNTHESIS FOR ALL TAXA 247 t of Threatened Threatened species ed as threatened, ed as threatened, ed as threatened gure does not provide does not provide gure All species cant gaps in its coverage for t may come to freshwater habitats, t may come to freshwater assessed at the 5% level was cance CrabsCrabsCrabs FishCrabs MolluscsFish OdonatesFish 0.48* PlantsFish 0.55*** 0.35Molluscs MolluscsMolluscs 0.18*** Odonates 0.12*** 0.48Odonates Odonates 0.19*** 0.71*** Plants 0.67*** Plants 0.67* Plants 0.09 0.14*** 0.68 0.16*** 0.72 0.13*** 0.86 0.24 0.08 -0.03 parks to freshwater systems. These results indicate that, systems. These results parks to freshwater although some benefi most often focuses areas management of protected environment. primarily on the terrestrial Using the distribution of species classifi congruent with are areas protected show that results of conservation concern only species freshwater across that 13.8% of found 4.90% of the total river network. We area outside the protected fall entirely species freshwater network, including 188 species classifi an indication of whether management within protected an indication of whether management within protected habitats. Nel of freshwater includes a consideration areas (2008) found that in South African National Parks, 50% to be considered of river systems within the parks were 72% of rivers outside parks in a poor state. However, impacted, suggesting some benefi considered were groups, such as bird, mammals and amphibians (Brooks mammals and amphibians (Brooks such as bird, groups, is also a bias in the location of protected 2004). There far at high altitude and are sites that are towards areas is of these areas as protection population centres, from upon them given the limited societal demands easier, two important questions (Joppa 2009). For these reasons, inclusive is the how species. Firstly, arise for freshwater network for inland waters of Africa, area protected current signifi there are and secondly, species? freshwater Database on the World Analysis based on data from (IUCN and WCMC 2010) suggests that Areas Protected continental Africa 10.21% of the river network across This is below the area. falls within an existing protected area within the protected 13.95% of the total land area network, and does not take into account the fact that as boundary markers to delineate the often used rivers are this fi Similarly, of reserves. borders Table 8.3. Spearman’s Rank Correlation between between Correlation Rank 8.3. Spearman’s Table species threatened and species richness total Signifi richness. account for spatial to correction using Dutieuls autocorrelation. ed as sh, molluscs, ts for others. sh or invertebrates. ed based on our ts for others. ts for others. 2007; Gaston 2008). With the exception et al. This lack of congruence presents ve taxonomic groups. cient allocation of resource for practical conservation cient allocation of resource of Ramsar Wetlands of International Importance, the of Ramsar Wetlands been targeted has rarely areas designation of protected species such as fi freshwater towards threatened according to the IUCN Red List (Table 8.3) for to the IUCN Red List (Table according threatened all fi a considerable challenge for conservation practitioners, as it indicates that investment in conservation for one group may not confer co-benefi ecosystems for freshwater areas 8.3.2 Protected scale drivers, broad to biodiversity from Faced with threats such as habitat loss and degradation, the emphasis of years from conservation action has shifted in recent the individual species towards one based on targeting scale broad that address areas establishment of protected of land for multiple species areas by managing threats (Langhammer identifi sites are Most commonly, knowledge of the conservation status of better-known work, and means that conservation planning for general work, and means that on information collected for a biodiversity can be based for globally assessed Surrogacy single taxonomic group. mammals and (for example, birds, taxonomic groups to be high, meaning that amphibians) is generally reported of one of conservation action based on the distribution will confer benefi these taxonomic groups Overall patterns richness for fi of species correlations strong odonates and plants exhibit reasonably between these 8.3), suggesting some surrogacy (Table scale geographic This is driven by broad taxonomic groups. but is lower than patterns in the tropics) (i.e., higher richness The between other taxonomic groups. has been reported groups between crabs and all other freshwater correlation of the perspective the strength a practical From is low. taxonomic indicates that although differing relationships patterns, they similar large-scale exhibit broadly groups for each other to enough surrogates not effective are the need for separate assessments. reduce by weak The need for separate assessments is highlighted between richness of species classifi relationships 8.3.1 Congruence between taxonomic groups between taxonomic 8.3.1 Congruence patterns and in species richness have considered we So far, the continent. biodiversity across to freshwater the threats of biodiversity freshwater how the rich now consider We conservation and how current Africa can be safeguarded strategies might be applied. conservation biology is how well data A central question in patterns in represents group collected for one taxonomic most allows the between groups High surrogacy another. effi 8.3 Conservation action action 8.3 Conservation 248 CHAPTER 8 | SYNTHESIS FOR ALL TAXA conservation measures are inplace. falling outsideaprotected area, inmanycases noknown classifi (Table 8.4).Thisisparticularlyworryingforthe30species List categoryfallingoutsideexisting protected areas. freshwater species. Number ofspeciesineachRed Table 8.4.Shortfallinconservationplanningfor the overlapbetweentwo. Figure 8.11.Distributionofprotected areas, areas identifi ols 21 24 147 1 215 0 1 DD 3 124 LC 16 12 NT 7 22 VU Odonata EN Mollusc CR Fish Crab Taxa ed asCritically Endangered where, inadditionto 000058 130015 ed aspotentialfreshwater KeyBiodiversityAreas, and birds andplants. for theconservation ofothertaxonomicgroups, suchas successfully applied toidentifythemostimportant sites for theidentifi be lost(Langhammer refers tothespatial optionsforconservationthatwill values willbelostinthefuture, andirreplaceability vulnerability refers to the likelihoodthatasite’s biodiversity the ‘vulnerability’and‘irreplaceability’ ofspecies.Here, using globallystandard criteriaandthresholds basedon are sitesofglobalsignifi protected area network(Langhammer tools developedtodetectandclosegapsintheexisting Key BiodiversityAreas (KBAs)are one of theprimary 8.3.3 KeyBiodiversityAreas forfreshwater species cation ofKBAshasbeendevelopedand cance forconservation, identifi et al . 2007).Themethodology et al. 2007).KBAs ed CHAPTER 8 | SYNTHESIS FOR ALL TAXA 249

sh sh cult, cult, (CR) (CR) t from from t nable boundary Giustia saidai Austroglanis barnardi ve odonates, 39 mollusc ve odonates, 39 mollusc cation for site designation is (CR) and sh, fi ) is known from a single pool in Angola single pool in Angola a ) is known from ned, for example, sh endemic to three tributaries of the Olifants sh endemic to three Iglica soussensis ed sites qualifying for designation under the AZE ed sites qualifying for designation under the AZE cation is based on three strict criteria. Firstly, the the strict criteria. Firstly, cation is based on three cant proportion of the species population occurs of the species population cant proportion Clarias cavernicola uctuations in the distribution of the species. immediate conservation action. The situation becomes is occupied the area for species where complicated more not as easily defi (EN) is a fi river whose range has extended at certain times into the main river channel. Unsustainable water abstraction has, to dry up, leading to caused this area years, in recent fl are known from single wells, where water abstraction is water abstraction single wells, where known from are leading to population decline. Species such as these are that would benefi small areas to incredibly restricted as river systems by their very nature are connected with connected with are as river systems by their very nature analysis, we landscape. In a preliminary the surrounding identifi of Critically Endangered criteria based on the occurrence species within a single sub-catchment. or Endangered Using these criteria, 134 fi and seven crabs would trigger AZE site designation. In a number of cases the justifi within which the character of the habitats, biological of the habitats, biological within which the character have more communities, and/or management issues with those in in common with each other than they do adjacent areas. date, the AZE analysis has focused on species groups To mammals, that have been globally assessed in entirety: amphibians, turtles and tortoises, iguanas, birds, corals, and conifers. The completion of crocodilians, species freshwater this assessment of many of Africa’s and exciting opportunity AZE with an interesting provides may qualify for to analyze at least the African species that species. AZE status, and identify AZE sites for those species, especially those in rivers, the For freshwater criterion makes designation of AZE sites diffi third cave catfi For example, the Critically Endangered clear. List, and if not safeguarded, the loss of these sites would sites would loss of these the if not safeguarded, List, and In species. or more of one extinction to the global lead based on new species analysis was updated 2010, the in 587 information, resulting and improved assessments CR or populations of 920 the last remaining sites that hold 141 sites, associated with Eighty two of these EN species. in the African continent. located are of these species, seen as subsets of KBAs, where AZE sites can be qualifi or Critically least one Endangered site must contain at the site is irreplaceable, species. Secondly, Endangered else on earth or a occurs nowhere as the species either signifi point in its life either permanently or at some there, the site must have a defi Thirdly, history. ( water abstraction ground measuring 18m by 2.5m, where the and collection for the aquarium trade threatens in northern of molluscs Africa a number species. Similarly, such as t t from an t from ed as potential freshwater ed as potential freshwater ed 561 sites around the world ed 561 sites around ection of the conservation status ection of the conservation rst identifi cation of freshwater KBAs. Some 2,305 sub- KBAs. Some 2,305 cation of freshwater KBAs found that 21.30% of the total area of these priority KBAs found that 21.30% of the total area network area sub-catchments falls within the protected 8.11). Uneven sizes and distributions of existing (Figure is far mean that coverage in some regions areas protected than in others. For example, existing comprehensive more in northern Zambia across areas Botswana, protected large good coverage of the and in south-east Angola provide such as the Okavango Delta. This contrasts with areas is little there Central African Republic and Senegal, where and potential areas overlap between existing protected and countries in western KBAs, freshwater Africa, where makes areas a highly fragmented network of protected complex. In all cases, understanding the relationship species to gain maximum benefi for freshwater however, Based on data collected during this assessment, the the assessment, during this collected on data Based Unit Biodiversity Freshwater Species Programme’s IUCN for criteria and thresholds site selection has developed the identifi selection Africa meet the continental across catchments of sites KBAs. The number potential freshwater criteria as is a clear refl qualifying (Christensen 1996) ecosystem management approach species as users of the water that legitimises freshwater and manages the catchment in a sustainable way. resource containing species of the highest conservation For areas can be applied in line the concern, strict protection more criteria (Dudley 2008). Area IUCN Protected areas protected Analysis of congruence between existing and sub-catchments identifi must management area, a protected inclusion within from of freshwater focus on the requirements include a greater ecosystems, with adequate consideration of longitudinal and lateral connectivity (see Chapter 9). Extinction – AZE sites 8.3.4 Alliance for Zero Extinction (AZE) includes membership The Alliance for Zero the world around of 70 non-governmental organizations of human-caused extinctions. dedicated to the prevention In 2005, AZE fi to that deserved immediate conservation action in order of 779 species. These sites hold the the extinction prevent (CR) of a Critically Endangered population last remaining to the IUCN Red species, according (EN) or Endangered of the continent’s freshwater habitats and associated freshwater of the continent’s species. delineated based on sub-catchment KBAs are Freshwater systems freshwater discussed, boundaries. As previously lowest point in the unique in that they occupy the are essential to consider biodiversity it is landscape, so to protect The form that this context. broader management in this dependent on vary considerably, management takes may of species present, and irreplaceability both the vulnerability basic level, alland the wider societal context. At the most KBAs would benefi sites qualifying as freshwater 250 CHAPTER 8 | SYNTHESIS FOR ALL TAXA assessment. Figure 8.12.ExistingAllianceforZero Extinctionsitesandthoseproposed asaresult ofthefreshwater 2020, then weneedtofocus conservation actionsurgently in partaimsto prevent theextinction ofspeciesthrough Target 11oftheConvention onBiologicalDiversity, which If weasaglobalcommunityare seriousaboutachieving and thisisshowninred withablueoutline. already designated2010 AZEsite(atBakossiMountains), one potentialfreshwater AZEtriggerspeciesfellwithinan AZE sitesidentifi AZE sitesinAfricared, withanadditional48potential their incorporation.Figure 8.12showsthe82offi AZE sitesinAfricawouldbeexpandedsignifi the freshwater speciesdatasuggestthatthenetworkof review expertsduringthenextAZEupdate, byexternal While notyetoffi ed forfreshwater speciesinblue.Only cially approved, giventheneedfor cantly upon cantly cial 2010 cial into new assessments. is anurgent priorityfor AZEtoincorporatethesenew data increasing thenumberofsitesby50%) inAfricaalone,it (nearly doublingthenumberoftrigger speciesinAfricaand in atleast136additionalspecies and48additionalsites in aprovisional draft analysis ofAZEqualifi to humans.Giventhatinclusion offreshwater biodiversity being, e.g.intermsofcontribution offreshwater services would provide disproportionate benefi Larsen when onlyconsideringAZEsitesforterrestrial species, species, butalsoprovide arangeofotherbenefi biodiversity conservationwillnotonlybenefi Additionally, safeguarding theseurgent sitesforfreshwater on thesesitesthat,iflost,wouldresult inextinctions. et al . (inprep) haveshownthatprotecting these sites ts tohumanwell- cation results cation t thetarget ts. Even ts. CHAPTER 8 | SYNTHESIS FOR ALL TAXA 251 shes and aquaticshes as well as providing income and trade. Overall, the most as well as providing fi the freshwater heavily utilized taxa are known species plants, with a total of 45% and 58% of all 8.13). Within the harvested (Figure harvested, respectively exploited for human consumption; species, 72% are consumed is of species that are the proportion however, 8.14). (Figure highly variable between taxonomic groups 8.4 Livelihoods dependence on 8.4 Livelihoods dependence on species freshwater species have been utilized by people throughout Freshwater to form an integral component in the and continue history, (see Chapter 1,livelihoods of millions of people in Africa for subsistence, upon directly relied Section 1.3). Species are Figure 8.14. Use of freshwater species by people in Africa 8.14. Use of freshwater Figure Figure 8.13. Percentage of assessed African freshwater species utilized by people. utilized by species freshwater African of assessed Percentage 8.13. Figure 252 CHAPTER 8 | SYNTHESIS FOR ALL TAXA © WORLDBANKPHOTOCOLLECTION afi Africa. Here tomanypeople'slivelihoodsin Fishing contributesgreatly crustaceans, molluscsandplantsplottedonlefthandaxis,fi Figure 8.15.Total reported harvest(intonnes)offreshwater speciesinAfricaoverlast60years.Freshwater supporting livelihoods.In Africa,thetotal annualfi waters are asimportanttodaythey everhavebeenfor Despite advances in,andexpansionof,agriculture, inland hisnetsinMozambique. repairs sherman shery low inAfrica, at2.1tonnes peryear, compared to21.4 Although fi income families (Millennium EcosystemAssessment 2005). for about70%to75%ofthetotal animalprotein forlow- 2004). InMalawi,forexample, inlandfi to ruralcommunitiesindeveloping countries(Kura where theyare oftentheonlysource ofprotein available species asaresource formillionsofpeoplewithinAfrica, These statisticsemphasisetheimportanceoffreshwater globally (Kura now seenasthefastestgrowing foodproduction sector fi and fl trade hasbeenmore recent formolluscsand crustaceans, been steadilyincreasing fordecades,expansionofthis fi accounting for25%oftheworld’s inlandwaterscapture and aquaticplantsismore than3.4milliontonnes, production forfreshwater fi rst timein2005(Figure 8.15).Aquaculture is,infact, sheries (FishStat2010).Whilstaquaculture offi shes ontheright.Source: FishStat2010. oral aquaculture production wasrecorded forthe shery production perperson is comparatively et al. 2004). shes, molluscs,crustaceans seis account sheries shes has shes et al.

CHAPTER 8 | SYNTHESIS FOR ALL TAXA 253 ts to sheries, with both sh species and for sheries statistics help to sh production in Africa. Uganda is among sh production shes are harvested, of which 81% are for harvested, of which 81% are shes are : Fishes are not only the most speciose of the not : Fishes are harvested (Figure 8.17); this includes species removed live 8.17); this includes species removed harvested (Figure no longer contributing for the aquarium trade, as they are species freshwater use of to the wild gene pool. Currently, impact on the long-term sustainability is having a direct for requirement of many wild populations in Africa. The is critical to sustainable management of these species of benefi their survival and for the continued provision many dependent people. units (that Monitoring of species use within management is, at the catchment and sub-catchment level) is important, to the species not only to identify potential threats concerned. In many cases, the Red List assessments of catch data, such as with provision could be improved fi has been used for some Lake Victoria fi where many species of sturgeon identify population trends. Fishes most but also the assessed here, groups freshwater a means utilized and depended upon. They provide as well as a means of income, sustenance, of direct Within Africa, 40% of known particularly for the rural poor. fi freshwater in the African countries are human consumption. Three in inland waters – production top 10 for global capture This highlights the particular Uganda, Nigeria and Tanzania. lakes for inland fi importance of the large Chad being ranked among the highest and Lakes Victoria fi for freshwater . et al sheries sheries sheries sheries. sheries and, as shers themselves, . 2010). et al ts to fi 2009; Dugan . 2004). By underestimating the scale . 2004). By underestimating et al et al. sheries employ approximately double that sheries employ approximately . 2004). Ready access to freshwater fi . 2004). Ready access to freshwater et al icting interests compete over use of water resources compete over use of water resources icting interests provides a real lifeline to many rural communities, acting a real provides local populations can as a ‘bank in the water’, where harvest species for rapid income needs, as well as acting as a ‘safety net’ for communities in times of agricultural (Béné hardship tonnes in Europe and a global average of 3.3 tonnes (FAO and a global average of 3.3 tonnes (FAO tonnes in Europe of 2008), this indicates the low level of industrialisation played by small-scale fi the trade and the key role the continent The majority of inland water catches across fi subsistence and recreational come from estimating that with the FAO unreported, such, go largely as likely to be two to four times as great actual catches are (Kura recorded are species, these resources of utilization of freshwater this poses is danger that great The under-valued. greatly when likely to be marginalised that small-scale needs are confl (Kura benefi In addition to the direct freshwater fi freshwater The fi number of people in the post-harvest sector. industry provides employment to the rural poor and poorly industry provides educated, and to women, with 55% of small-scale fi workers in developing countries being female (Dugan 2010). species utilized in Africa The vast majority of freshwater of unknown origin (Figure the wild, or are taken from are is lethal, as the whole organism 8.16). Use is predominantly Figure 8.16. Source of utilized freshwater species in Africa. species freshwater of utilized Source 8.16. Figure 254 CHAPTER 8 | SYNTHESIS FOR ALL TAXA *nei – Not Elsewhere Included – Species groups where individualcatch datawasnot collected atthespecies level. Included –Speciesgroups where *nei –Not Elsewhere Table 8.5.Top fi Figure 8.17.Formofremoval ofutilizedfreshwater speciesinAfrica. stocks; indeedthemajorityofAfrica’s fi are gettinglarger, thisisnotanindicationofhealthy fi Although theindustryisincreasing andoverallcatches Nigeria (Table 8.5). gariepinus pike ( niloticus mullet ( The fi globally (afterLatinAmericaandtheNearEast;FAO 2008). 2006, Africashowedthethird largest growth inaquaculture is becomingagrowing industry, andbetween1970 catches are largely from naturalsystems,butaquaculture highest freshwater fi the smallestofAfrica’s nations,yetcurrently claimsthe Ghana 5,594 Nile Tilapia, NorthAfrican catfi NileTilapia, sh, Africanbonytongue Three spotted tilapia(Oreochromis NileTilapia, andersonii),Longfi n tilapia Torpedo-shaped catfi zillii),Common (Tilapia Red-belliedTilapia shes (nei),NileTilapia, 5,594 CommonCarp(Cyprinuscarpiocarpio),Torpedo- FlatheadGrey Mullet,Nile Tilapia, 5,640 KafuePike,NorthAfrican catfi sh, Cyprinids(nei),Torpedo-shaped catfi shes (nei),Nile Ghana 52,250 98,833 Zambia Uganda 143,207 Egypt Nigeria Country ve mostcultivatedspeciesinAfricaare: fl Hepsetus odoe Mugil cephalus ), ‘torpedo-shaped’catfi ). Byfarthelargest aquaculture production isin ve Africancountriesrankedbyinlandaquaculture production in2008(source: FishStat2010). rdcin()Top cultured species production (t) 2008 Aquaculture sh catchonthecontinent.These ) andNorthAfricancatfi ), Niletilapia( shes ( shes Oreochromis niloticus Oreochromis Clarias sheries are either (Oreochromis macrochir), rendalli), Redbreast Commoncarp tilapia(Tilapia Carp, NilePerch shaped catfi shes (nei),European seabass Perch (Latesniloticus) athead grey spp.), Kafue s ( sh Clarias sh in importance(forexample, in Nileperch catches,othernativespeciesare escalating fi and and evenPossiblyExtinct(suchas perch, and73%oftheseare nowCriticallyEndangered out ofthe74threatened cichlidsare threatened byNile has created amulti-milliondollarindustry. However, 64 In LakeVictoria forexample,theintroduction ofNileperch while dramaticdeclinesofnativespeciesgoun-noticed. ecosystem maybemaskedbyanoverallincrease incatch species havebeenintroduced, theoverallimpactson fi sometimes easytosee,asillustratedbythecrashin fully- orover-exploited (Brainerd 1995).Theeffects are particularly forlocalsubsistence.Theseshiftsinreliance shed speciesinLakeVictoria, andamore recent decline shery production inLakeChadthe1980s.Where H. fl avipinnis ). Withmassivedeclinesintraditionally Haplochromis laparogramma Haplochromis dentex )

CHAPTER 8 | SYNTHESIS FOR ALL TAXA 255 shing, Lamprologus . 1997), whereby . 1997), whereby spp.) are the most the are spp.) shing practises still et al is one of three chambo is one of three (EN) in Lake Mai-N’dombe, (EN) in Lake Mai-N’dombe, shing result in high by-catch, shing result Oreochromis © WILLIAM DARWALL (VU) in Lake Tana, and (VU) in Lake Tana, shes and, as a result, some species are species are some as a result, and, shes Oreochromis karongae (EN) in Lake Tumba. Some fi (EN) in Lake Tumba. process in the to result shing has been demonstrated down the food web’ (Pauly shing shing activities, particularly through the use of nets with activities, particularly through shing with as much as a 70% decline in catches within a 10-year with as much as a 70% assessed as Endangered. period, and it is now non-targeted through threatened Other species are fi example, mosquito nets). These small mesh sizes (for of fi indiscriminate methods impact on some and may have a severe especially of fry, in smaller lakes. Examples include species, particularly Alestopetersius nigropterus Garra regressus tumbanus exploited as levels are species occupying lower trophic In Lake Malawi, ‘chambo’ ( ‘chambo’ Malawi, In Lake food fi valuable become they have where to the point being overexploited threatened. for which is targeted to Lake Malawi, species endemic of this Populations as well as commerce. local subsistence of overfi as a result in the 1990s species collapsed impact, reaching used in parts of Africa have an even wider poisons. Such such as the widely illegal practise of using indiscriminate and may even degrade the methods are underlying habitat itself. Overfi of ‘fi shed, and highly shed, © GEORGE TURNER (EN), a heavily overfi ed for human uses, and long-term changes between different species highlight the importance of between different can adapt to maintaining a rich diversity of species that The ability of ecosystems throughout changing pressures. important as more Africa to adapt is becoming increasingly become modifi due to climate change (see section 8.6). projected are Oreochromis karongae Oreochromis valued, species endemic to Lake Malawi. Fishing provides an important food source for many people living in the Okavango Delta. Fishing provides an important food source for many people living in the Okavango Delta. 256 CHAPTER 8 | SYNTHESIS FOR ALL TAXA of reasons, predominantly As forfood, orasadornments. bivalves andgastropods havebeenexploited foranumber source oflivelihoodthroughout Africa.Formillennia,both Molluscs: local livelihoods(BicknellandChin 2007). conservation (forinstanceinGuyana), aswellproviding serve asadirect incentive forhabitatpreservation and for wild-harvestedaquariumspecieshasbeenproven to habitat degradation.Establishingasustainablescheme usually threatened duetotheirrestricted range andlocal currently thoughtamajorthreat. Thesespecies are more although over-harvesting fortheaquariumtradeisnot targeted fortheaquariumtradeinAfricaare threatened, the wild.Eightytwopercentofspeciesnotedasbeing (Tlusty 2002),yetlarge numbersare stillharvestedfrom freshwater fi being collected.Itisbelievedthatover90%ofornamental aquarium trade,with8%offreshwater fi The secondmostcommonutilizationoffi 2004). species, asweseetoday(Barrett, LockerandRobert on freshwater speciestowards exploitationofmarine around athousandyearsagoledtotheshiftfrom reliance that collapsesinfreshwater fi illustrated byrecent archaeological evidencesuggesting impacts canhaveprofound socialconsequences,as options foradaptationtoachangingenvironment. Such species alsoreduces geneticdiversity, soreducing the impact onprimaryproductivity (Allan cycling suchthatthelossofspeciescanhaveasignifi system. Forexample,fi can haveaprofound infl et al some cases,collapseofthefi increase inthetotalcatch,followedbystagnationand, unprofi those speciesattheapexare lost,orbecomecommercially © WILLIAMDARWALL forms partofthefi forms ( The yellow-bellybream . 2005).Theresulting changeinthefoodwebstructure table astheirnumbersdecline.Thisleadstoaninitial Mollusc collectionhastraditionally beena shes canbebred successfullyincaptivity shing catchintheOkavangoDelta. sh playanimportantrole innutrient Serranochromis robustus jallae uence onthefunctioningof shery (Pauly sh stocksacross Europe et al. et al 2005).Lossof shes inAfrica sh isforthe . 1997;Allan ) (LC), cant to focus primarilyonfreshwater fi often overlooked, asoffi Crabs affi ( only three species were recorded asbeingcollected numbers ofmosquitoesindomestic watertanks.InAfrica, they are specifi dragonfl that are notregularly usedbypeople.It isknownthat Odonata M. subgraëllsiana mourebeyensis mourebeyensis Melanopsis letourneuxi where speciesdirectly impactedbycollecting include from thisactivityareAfrica, particularlyhighin northern range thisharvestingrepresents amajorthreat. Threats practise formanyspecies,somethatare restricted in as jewelleryanddecoration.Althoughthisisacommon Mollusc shellsandpearlsare frequently harvestedforuse impact onspeciessurvival. and suchactivitiesare notthoughttobehavingamajor enthusiasts. However, thismarketremains relatively small, sought afterandcanfetchahighprice,particularlyamong species isspecimencollection.Rarer shellsmaybegreatly The formofusecommontothegreatest numberof (Pople 1966). important source ofmeat,producing upto112kg/ha/yr elsewhere withinitsnaturalrange,providing alocally the world.However, farmingofthisspeciescontinues the 1960sofLakeVolta, thelargest reservoir byarea in of the 1960s(Pople1966).Thesefi bringing inanannualharvestof4,000to7,000tonnesby been transportedintoupstream fi the Volta Riveraloneitisbelievedthatyoungclamshave Cross, ItuandSanagaRivers.Itiseasytomaintain, and in Africa, where itisfoundinthelowerreaches oftheVolta, Egeria radiata harvested from thewild,butare alsofarmed.Forexample bait (e.g., are alsocommonlycollectedforanimalfeed,orasfi (e.g., Based onvolume,theprimaryuseofmolluscsisforfood rural communities. of thefewincome-generatingoptionsopentowomenin in inlandwatersisbywomenandchildren andcanbe one industry forwomen,asthemajorityofmollusccollection decoration orjewellery. Itisaparticularlyimportant in comparisontofi rural communities,molluscsoftenfetchhighmarketprices well asbeinganimportantformofsustenancetosome Aethiothemis bequaerti nis E. radiata ), butitisnotknownforwhatpurpose. Pila occidentalis : Theimportance ofcrabsasasource offoodis y nymphsare usedasfi : Odonatesare theonlyfreshwater taxaassessed Chambardia wahlbergi) ) havediminishedduetotheconstructionin isafreshwater clamendemictowestern cally usedaspest-control to reduce the (EN), (VU). shes, andevenmore soasitemsof M. saharica or , (EN), cial fi cial Aciagrion africanum Mutela zambesiensi sheries statisticsmaytend (CR), sheries (andpopulations M. magnifi shing bait,andinIndia . Molluscsare notonly sh species. However, sheries forcenturies, M. scalaris ca and ). Molluscs (EN), (EN)and Aeshna shing M. CHAPTER 8 | SYNTHESIS FOR ALL TAXA 257 Xyris used for , is used in bre, bre, and Cyperus papyrus Ambrosia maritime Lasimorpha senegalensis © ROGER BILLS/SAIAB crabs; and and odonates shes, molluscs, for food or as a medicine. Collecting water lilies in Mozambique for human consumption. human consumption and decoration, of fi as a structural material and a source anceps use of plants is for medicinal The most widespread have a vital purposes, for people and livestock. Plants as medicines, particularly in rural communities where role may be extremely access to other forms of healthcare limited. For example, and to remedy traditional medicine to expel kidney stones asthma, and even bilharzia. troubles, renal group the only freshwater Aquatic plants represent 8.17). to have non-lethal harvesting (Figure reported cultivation Plants also have the highest level of captive wild 8.16). For the most part, therefore, (see Figure by overharvesting, not threatened populations are and restricted exceptions for scarce are although there region are varied, with are region

, for Typa Typa sh (Neiland ); medicinal . Juncus rigidus ). In many cases, Cyperus papyrus spp Persicaria bistorta 2011). Uses in this

Butomus umbellatus, Typha spp., Cyperus Typha , (LC), , ); or for cultural reasons ); or for cultural reasons cinalis Ceratophyllum demersum Nymphaea species for water gardens, species for water gardens, (LC) (Ghogue some Genista ancistrocarpa ornamentation and crafting household . 2009). Due to their size and biomass, . 2009). Due to their size and biomass,

, ., Gratiola offi Bacopa monnieri : Plants are commonly utilized throughout commonly utilized throughout : Plants are , Nympahea et al spp Phragmites australis , Lasimorpha senegalensis Xyris anceps ); decoration (e.g., as an ornament); food (e.g., Trapa natans Trapa purposes (e.g., use in aquaria, (e.g., indirect evidence from infection rates of paragonimiasis evidence from indirect (a food borne and other diseases for which zoonosis) intermediate hosts suggests that consumption crabs are the world throughout crabs is widespread of freshwater (Cumberlidge of protein, an important source crabs can represent suggesting that for poor people living far with research on may be a heavier reliance there water resources from on fi crabs and other aquatic animals than and Bene 2008). Aquatic plants is a rapidly taxa, there the world. As with other freshwater and, in 2006, market for plants in aquaculture growing was 15.1 million tonnes with global aquatic plant production all virtually 2008). However, a value of USD 7.2 billion (FAO 2008), and very Asian markets (FAO of this comes from in Africa. Data of plants little is known about aquaculture still plants are for Africa indicates that utilized collated here only 8% of the wild, with harvested from predominantly 8.16). used species known to be cultivated (see Figure collected for a In contrast to the fauna, aquatic plants are are 8.18). The uses recorded (Figure wide range of reasons and include: structural materials (e.g., and varied numerous Juncus spp papyrus latifola Aponogeton Mentha cervina species have multiple uses. For example, is used medicinally to treat swellings and gout, has edible is used medicinally to treat shoots, is used for used to make the stems are items, and in Tibesti, Niger, brushes. In central Africa, the most heavily used freshwater plants are (LC) and Figure 8.18. Utilization of freshwater species in Africa: a) Freshwater fi Freshwater Africa: a) in species of freshwater Utilization 8.18. Figure plants. b) Aquatic 258 CHAPTER 8 | SYNTHESIS FOR ALL TAXA considered threatened. collection and heavy grazingpressure bylivestock,are used byhumans and,through thecombined pressures of azureus number ofspeciesfrom thefamily, including pondweed isnotthreatened through over-exploitation, a fl Cape dish‘waterblommetjiebredie’ hasCapepondweed commercial purposes(Sieben2009),andthetraditional South Africa.Itsfl Capein as Capepondweed,isendemictothewestern Africa. southern well knownfortheiredibletubersandare consumedin due toitsmedicinalproperties. endemic toMorocco, isthreatened by overharvesting populations. Forexample, Figure 8.19.Sub-catchmentswithfi owers as itsmainconstituent.AlthoughtheCape (VU)and Aponogeton distachyo Aponogeton bogneri owers are eatenandcultivatedfor Mentha gattefossei Aponogeton spp ve ormore DDfi , commonlyknown (EN)are similarly Aponogeton , aperennial . are sh speciesandatleastonethreatened species. species coincide withareas thoughttobe threatened might fi lack ofinformation can,inmanycases,befi the broader ecology of theregion beingexamined.This ecosystems, infersasignifi to represent awiderangeoftrophic levels withinfreshwater knowledge forspeciesinthese particular groups, selected species havebeenhighlightedin chapters3to7.Thelackof where found.Severalregions withhighnumbersofDD the impactofpotentialthreats withinthesub-catchments there isinsuffi Species are oftenrecorded asDataDefi 8.5.1 RegionsofDataDefi 8.5 Areas forfurtherwork eld survey. Sub-catchmentswhere high numbersofDD cient informationontheirdistributionsand cient andthreatened species cant paucityofknowledgeon cient (DD)when le through lled CHAPTER 8 | SYNTHESIS FOR ALL TAXA 259 uent of the Kotto River ra River in the Upper Lualaba ecoregion; Lake Mweru ecoregion; ra River in the Upper Lualaba (Figure 8.19), include: the Konkoure and Rokel basins in the Konkoure 8.19), include: (Figure the upper part of the Cross the Upper Guinea province; River; the Nyong, Kribi, Ivindo, Kouilou/Niari in the Lower much of the lower and middle Congo Guinea Province; up to the tributary of the Ruki River at Mbandaka; the Ubangi between Bangui and the affl Moboye; the Upper Congo Rapids; the from upstream Lufi and the Kafubu River in the Bwengulu-Mweru ecoregions; parts of the Lovua, Lulua and upper Kasai basins in the the Cuanza River in Angola; Lake Tana Kasai ecoregion; (and basins immediately to in Ethiopia; and Lakes Victoria and Malawi. the north-east of the lake), Tanganyika, For odonates, sub-catchments with both high levels of DD 8.20) species (Figure species and at least one threatened ciency ciency are sh species, eld survey. eld survey. eld survey. An analysis of eld survey. es a number of such priority ve or more DD fi ve or more ciency for two taxonomic groups, ciency for two taxonomic groups, ciency, with the exception of those ciency, ve or more DD species in a single sub- ve or more shes, fi shes and odonates, identifi be considered high priorities for fi high be considered catchments. Sub-catchments with a to other relative of DD species (e.g., > 25% of the total high percentage also taken to represent were number of species present) high levels of data defi sub-catchments with very low total numbers of species. Sub-catchments with fi species was known to be threatened at least one where the patterns of data defi a high level of data defi catchment represents fi serving preliminary, are here presented The results areas. patterns as a test of concept where of data defi fi for future used to identify priority areas For fi Figure 8.20. Sub-catchments with both high levels of DD odonate species and at least one threatened species threatened least one and at species of DD odonate levels both high with Sub-catchments 8.20. Figure 260 CHAPTER 8 | SYNTHESIS FOR ALL TAXA identifi Lakes Victoria andTanganyika, coincided withthosealso Several ofthese sub-catchments,suchasthose around north toLakeTanganyika inthesouth;andLakeVictoria. the sub-catchmentsextending from LakeAlbertinthe Lac Tsolo inthenorthandalongborder withCongo; Gabonfromthe east;south-eastern Diengainthewestto basins tothecitiesofBamenda, Yaounde, andBafi Cameroon,Liberia; muchofsouthern includingallcoastal Nuon, Cestos,Dugbe,andSehnkwehnsub-catchmentsin sub-catchment inGuinea;theDu,Nianda,SaintJohn, and Bagbesub-catchmentsinSierraLeone;theFaliko include: theMabole,Seli,Pampana,Rokel,Taia, Sewa, richness. Figure 8.21.Sub-catchmentswithhighnumbers of DDfi ed for fi edfor shes. a in a sh speciesadjacenttoregions withlowoverallspecies numbers ofspecies(i.e., lessthanfour speciespersub- species are adjacenttoregions withrelatively low overall Several sub-catchments withhighnumbers ofDD priority forfuture fi endemic totheselakes.Areas suchasthesemustbea are threatened, butalsobecausemostofthespeciesare particularly disturbing,notjust becausethesehabitats DD speciesinLakeTana andtheAfricanGreat Lakesis other speciespresent. Thepresence ofhigh numbersof basic ecologicalinformationforahighproportion ofthe threats toatleastonespecies,yetthere is alackofthe The regions identifi eld survey. eld ed represent placesthere are known CHAPTER 8 | SYNTHESIS FOR ALL TAXA 261 shes in the sh species and are species and are sh sh species. One might, sh species. One might, eld survey, this would imply that this would imply that eld survey, following additional fi there to the zones where restricted truly the species are known threats. are DD species within sub-catchments surrounded Finally, by sub-catchments with high numbers of threatened as likely 8.23) might be considered species (Figure example, the Nyong and Kribi basins For threatened. the Ivindo and Bouniandje basins in and in Cameroon, Gabon have high numbers of DD fi by sub-catchments with almost completely surrounded fi high numbers of threatened that the fi predict reasonably therefore, cient cient shes, which is the median value for all shes, which is the median value for all catchment for fi found in these adjacent catchments also catchments are lower and may represent apparently are threats where not found if the DD species are safe havens. Conversely, the sub-catchments in Africa) and no threatened species the sub-catchments in Africa) and no threatened (e.g., parts of the lower Congo and upper Ubangi in basin in eastern Africa) Figure central Africa; the Tana sub-catchments in for odonates, are 8.21. Similarly, north-eastern Namibia and north-western Botswana in numbers of species low recorded 8.22). The (Figure of course, be an these neighbouring catchments may, artefact of limited sampling. If this is the case, then it is in the data defi possible that the species present Figure 8.22. Sub-catchments with high numbers of DD odonate species adjacent to regions with low overall low overall with to regions adjacent species DD odonate of high numbers with Sub-catchments 8.22. Figure richness. species 262 CHAPTER 8 | SYNTHESIS FOR ALL TAXA above, will beespecially relevant and usefulfor of datadefi Information on patterns 8.5.2 RapidAssessmentPrograms (RAPs) threatened odonatespecies. species adjacenttobasins withhighnumbersof Lac deBamendjing,havehigh numbersofDDodonate catchments, aswellthesub-catchmentssurrounding Nkam, Woun, Dibamba,Noun,andtheCross sub- slightly intoNigeria,includingtheMungo,Wouri, Nita, coastalCameroonlocated inwestern andextending investigated asapriority. Similarly, sub-catchments threatened themselves;thesespeciesshouldbefurther Nyong, Kribi,IvindoandBouniandjeare signifi numbers ofthreatened species. Figure 8.23.Sub-catchmentswithhighnumbersof DDfi ciency, as discussed cantly sh speciessurrounded bysub-catchmentswithhigh conservation challenges. data canbeavailable tomeettheneeds ofurgent that biodiversity dataare collected quicklysothatthe This techniqueofrapidbiodiversity assessmentensures of thehealthecosystem and itsconservationvalue. focused ontaxonomicgroups thatcangiveanindication RAP fi biological valueoflittle-known andunexplored areas. expert scientiststoconductfi andhost-country RAP assemblesteamsofinternational needed tomakequickbutsoundconservationdecisions. in 1990toaddress thelackofbiologicalinformation the fi (RAP) anditspartnersinthevariouscountrieswhere Conservation International’s RapidAssessmentProgram eld programs are implemented.RAPwascreated eld surveysgenerallylastthree tofour weeks, rst-cut assessmentsofthe CHAPTER 8 | SYNTHESIS FOR ALL TAXA 263 et rst shes . 2003; et al Afrithelphusa ed Forests in ed Forests rst time since its shermen and sport sh species and one rst time in these areas, and rst time in these areas, 2008). The AquaRAP data are 2008). The AquaRAP data are et al. sh species were recorded, including recorded, were sh species sp. nov.; Dijkstra 2007b). sp. nov.; icts between local fi . 2003). These AquaRAP surveys provided surveys provided . 2003). These AquaRAP c understanding of species distributions, distributions, of species c understanding , which was recorded for the fi , which was recorded et al Eluthemis 2005). shermen in the delta. Data on the shrimp fauna from in the delta. Data on the shrimp fauna from shermen many ways to further conservation efforts and to advance to advance and efforts conservation ways to further many our scientifi surveys have provided RAP and biology. habitats, the distribution data for to update species information science, species new to List, have discovered Red IUCN’s time, and not seen in a long species have re-discovered records and new country range extensions have recorded for many species. two surveys of the Okavango delta AquaRAP carried out high water in 2000 and the second in Botswana, one in 2003. One fi during low water in aquatic Heteroptera (Belastomatidae) new to science (Belastomatidae) aquatic Heteroptera along with several survey, during the 2000 recorded were the Okavango delta (Appleton for new records Tweddle management and diversity data for essential freshwater a process monitoring of the Okavango Delta and catalyzed confl for resolving fi being used to assess health risks in the 2000 survey are the delta (Appleton to nominate being used to support a proposal currently Heritage Site. the Okavango delta as a World diversity was also surveyed along with terrestrial Freshwater Liberia, Guinea, taxa during RAP surveys in Côte d’Ivoire, Africa, and in D. R. Congo in Central and Ghana in West Africa. In Haute Dodo and Cavally Classifi 33 fi Côte d’Ivoire, for the fi nine species recorded cavalliensis) endemic to the one species (Chromidotilapia by IUCN) (Gourène Cavally River (listed as Vulnerable al. of western Guinea, the RAP team In the Boké Prefecture purple marsh crab, the Endangered recorded monodosus in 1947 (Cumberlidge 2006). This species original record cultivated land, so little is still known was collected from One species of about its natural habitat and biology. fi in Boké for the was also recorded shrimp freshwater from time in Guinea, extending its known range westward Sierra Leone (Cumberlidge 2006). during the RAP survey of Lokutu sampled Odonata were with recorded, in D. R. Congo. A total of 86 species were 62 (72%) Guineo-Congolian species, 13 of which have not species been found outside of the Congo Basin, and three also likely new to science (Dijkstra 2007a). Odonata were in Liberia in 2005. RAP national forests sampled in three 93 species, with one species new to documented results science ( surveys in Ghana included assessment of RAP Two fi and macroinvertebrates taxa. Freshwater freshwater the Ajenjua Bepo and Mamang River surveyed in were shes, ow, ow, and ed above might © JENS KIPPING rst-cut assessment of cally target freshwater freshwater cally target taxonomic ed to the most appropriate ‘AquaRAP’ surveys specifi catchment when ecosystems, focusing on the entire organisms possible, to include all the habitats that aquatic teams typically use during their life cycles. AquaRAP internationallocal institutions include scientists from and experts on the taxonomy of aquatic plants, fi who are and insects), aquatic invertebrates (molluscs, crustaceans also a key are and plankton. Specialists in water quality physical data on the relevant part of the team, to provide fl parameters (e.g., dissolved oxygen, turbidity, level (genus or species) to obtain a fi nutrient levels) to help explain patterns in the species goal of AquaRAP is to assess the data. The overarching based biological and conservation value of a catchment, of the habitats; (2) a preliminary on: (1) the heterogeneity that characterize each of these survey of the organisms of habitats, and habitats; and (3) the overall intactness resources their capacity to support important biological made to Field collections are and ecological processes. minimized. but the collections are identify the organisms, identifi Specimens are diversity and species composition. critical data employed in AquaRAP surveys have provided many cases for assessments of species Red List status in It is, therefore, biodiversity. freshwater this study of Africa’s identifi hoped that the priority survey areas Africa. The value of stimulate further surveys throughout AquaRAP surveys as input to the species assessments is an overview of the assessments demonstrated through conducted in Africa to date. Since 1990, Conservation International has conducted 13 AquaRAP surveys that have focused on the aquatic catchment. Several other the entire ecosystems through focus also included RAP surveys that had a terrestrial aquatic taxa. Eight RAP surveys conducted in Africa surveyed aquatic taxa. The RAP data have been used in Field survey the Okavango Delta channel. in 264 CHAPTER 8 | SYNTHESIS FOR ALL TAXA result ofintensifying human modifi phenology) tonewconditions has becomereduced asa to species(suchaschanges in range,behaviour, and shifts, and(b)thesetofecological responses available that (a)humansare theprimarydriverofcurrent climate climate changeisnovel,notinitspaceofbut evolutionary timescales.However, thecurrent periodof impacts onspeciesandecosystems,particularlyover climate changeisnotnecessarilyasource ofnegative for aslongtheAfricancontinenthasexisted.Byitself, African ecosystemshaveexperiencedclimatechange 8.6.1 to freshwater species–climatechange. look aheadandconsideroneofthemostseriousthreats habitats forfuture generations.Inthisfi greater investmentinunderstandingandmanagingthese across thecontinent.Thismakesacompellingcasefor that freshwater specieshaveforlivelihoodsofpeople what hasemerged through thisstudyistheimportance conservation ofspeciesshouldbeagoalinitsownright, and improve ourknowledgeofspecies.Althoughthe and somepracticalstepsthatcanbetakentoconserve in Africanfreshwaters, thethreats thatthisdiversityfaces, In thischapter, wehaveconsidered therichdiversityoflife 8.6 Thefuture ofAfricanfreshwaters gaps identifi assessments across Africatofi in thisreport willencourageanewseriesofAquaRAP In conclusion,itishopedthatthefi montane environments (Dijkstra2007c). (regionally classifi addition, thisspeciesisaregionally threatened dragonfl were usedinthespeciesdescription(Dijkstra2007c). In it hadnotbeendescribedatthetime,soRAPdata Atoconeura luxata of dragonfl from Côted’Ivoire (Abban2007).Amongthe72species were documented,includingonepreviously knownonly the AtewaForest Reserve,inGhana.Nineteenfi Also in2006,aRAPteamsurveyedOdonataandfi time inGhana. pobeguini Samman andAmakye2008).Onefi were typicalofdisturbedordeforested areas (Abban2008; Forest Reservesin2006.Manyspeciesofbothgroups ¹ will refer toCCA. anticipated climatechange impacts;‘adaptation’inthisusageisnot directly connectedtostandard ecologicalandevolutionar Climate changeadaptation (CCA)isatermdatingbacktoabout1990, borrowed from theclimate changepolicycommunitytodescr conservation: seeingthefuture through thepast Climate changeandAfricanfreshwater (LC) ies anddamselfl ed. , atMamangRiverwasrecorded forthefi (LC)wasespeciallysignifi ed asvulnerable),foundusuallyin ies recorded, therecord of ll themanyinformation cation of ecosystems s species, sh nig presented ndings nal section,we cant because cant sh species shes in shes Barbus rst y fl Congo basincarriesabout30%ofAfrica'ssurfacewater atsunset,Yaekela, returning Fishermen D.R.Congo.The as anewclimateregime beginstoemerge. on enablingorrestoring theseclimate-adaptivecapacities changes, andconservationefforts shouldprobably focus shown resilience inthe faceofdramaticeco-hydrological climate historysuggeststhatecosystemsandspecieshave and Wickel2009;Wilby called ‘climatechangeadaptation’framework(Matthews may represent abreak withcurrent practicesunderaso- informed conservationofAfricanfreshwater ecosystems of ongoingclimaticshifts–andinsomecases,climate- freshwater speciesandecosystemsmusttakeaccount and freshwater species.Conservationefforts forAfrican that willbesustainableforAfrica’s developingeconomies may beusefulguidestofuture freshwater management and howpastcurrent ecosystemandspeciesresponses projected impactsrelated toanthropogenic climatechange; of Africanfreshwater systems;observed,emerging, and This sectionwillbriefl al. change inducedeco-hydrological impacts(LeQuesne ability ofecosystemsandspeciestoadjustclimate that manyofthesemodifi industrial, andurbaninfl water use,and(often)limitedtreatment ofagricultural, infrastructure, direct modifi has increasingly involvedthedevelopmentofwater 2010). Economicdevelopmentforthepasttwocenturies recent decades(Dudgeon have seendramaticpopulationandspecieslossesin (Parmesan 2006).Freshwater ecosystemsinparticular ow. 2010). © KLAAS-DOUWEDIJKSTRA y describe:therecent climatichistory et al y usesoftheterm,and adaptationinthissection ows. Aconsensusisemerging . 2010)¹.However, Africa’s recent cation ofchannels,intensifying et al. cations reduce theinherent ibe efforts torespond torealized and 2005;Vörösmarty et al. et

CHAPTER 8 | SYNTHESIS FOR ALL TAXA 265 et ). Other uctuations uctuations in press uenced by their . 2008). Kröpelin uence of recent uence of recent . 2003; Marchant to ow unimpeded . 2006). Deposition et al et al . 2009). Water quality . 2009). Water et al et al uxes (Freeman 2007). In uxes (Freeman . 2008). uxes from precipitation and precipitation uxes from et al cation processes (Marchant and (Marchant cation processes . 2008; Dupont et al A short history of African eco-hydrological A short history of African eco-hydrological dynamism: the past 100,000 years . (2008), for instance, document the gradual shift in. (2008), for instance, document the gradual . 2007; Conway over the last 20 million years, and none fl and none million years, last 20 over the lakes 1997). African (Lévêque continent of the the interior fl but large synchronous relatively tend to show in level. Over the 20th century, many large lakes have lakes have many large the 20th century, in level. Over to 10m (Nicholson 2000). varied by 5 by ‘green Africa is dominated than 70% of more Additionally, fl water’, or atmospheric-water as opposed to ‘blue’, or terrestrial evapotranspiration, water fl surface and aquifer patterns, seasonal precipitation combination with highly exposed to regularly ecosystems are many freshwater most of the continent, high Across weather extremes. variability is not interdecadal, interannual, and even mean of 20 to 40% from uncommon, with variations instrumental record often found in the annual precipitation (Nicholson 2000; Conway 8.6.3 freshwater characteristics of Africa’s The eco-hydrological extensive shifts over the past systems have undergone of the Holocene100,000 years, and even since the onset 12,000 years BP; Gasse 2000; Issar 2003). Given (around deeply infl that ecosystems and species are the infl evolutionary and ecological history, conditions may serve as a useful guide to eco-hydrological impacts. future in African climate and hydrology of trends Recent reviews conditions over the past 100,000 years suggest that current of the to the rest compared dry, relatively in Africa are Holocene (e.g., Gasse 2000; Holmgren 2006; Weijers and Hooghiemstra 2004; Kiage and Liu et al northern a relatively Africa over the past 10,000 years, from arid state within the past to its current Sahara’ wet ‘green large, ecosystems evolving from 3,000 years, with freshwater lacustrine and wetland systems topermanent freshwater temporary and/or saline (or hypersaline) systems (Nicholson Hooghiemstra 2004). Similar patterns and 2000; Marchant paleoecological in Lake Malawi, where seen to the south are studies document rapid transitions between ‘dilute’ states comparable to the contemporary system to highly alkaline water in or drops of increases and saline conditions as a result (Stone of metres level of dozens to hundreds Hooghiemstra 2004; Marshall and with major shifts in productivity has varied accordingly, species assemblages (e.g., Cohen show that rivers such as the Nile and Congo are records al African Rift valley lakes have also expanded and contracted to shifting directly millennia, responding dramatically in recent (Gasse 2000). Several studies show regimes precipitation that shifts in monsoon seasonality have been associated with changes in lake stratifi . ow, ow, et al . 2008). et al . 2005; Abell . 2005; Abell et al © ROBERT SCHELLY © ROBERT ve climatic zones exist across the ve climatic zones exist across owing only seasonally (Lundberg owing only seasonally (Lundberg . 1995; Thieme . 1995; Thieme et al Africa’s river and lake basins are among the oldest in the among the oldest and lake basins are river Africa’s the Miocene originating before world, with most basins 1981). 24 million years BP; Beadle (approximately fi at least Currently, Mediterranean, subtropical, tropical, continent: equatorial, pattern precipitation is one of and mountain. The general the equator in the Congo Basin and highest rainfall near less at higher with progressively along the Gulf of Guinea, rainfall and low evaporation, the latitudes. Due to its high surface fl about 30% of Africa’s Congo Basin carries African rivers is 2000). Another distinctive characteristic of rapids and waterfalls that they generally have many more tectonic activity than other rivers in the world because of though accounting for only about 13% of the surface area the surface area though accounting for only about 13% of 1995). Arid, semi-arid, and dry sub- of the continent (FAO and surface area cover about 43% of Africa’s humid areas the north and south (Bjørke 2002). deserts in include large less than 9km rivers are of Africa’s than 90 percent More long, with many fl 8.6.2 African waters: special qualities and challenges qualities special African waters: 8.6.2 a number possess ecosystems freshwater Extant African and ecological characteristics of macro-hydrological continents systems on other freshwater distinct from (Cushing Foulakari Falls, D. R. Congo. African rivers generally have more rapids and waterfalls than other rivers in the world, due to the history of tectonic activity across the continent over the last 20 million years. 266 CHAPTER 8 | SYNTHESIS FOR ALL TAXA changes inproductivity. affect stratifi lakes, suchasLake Victoria, may of regime Changes inthethermal cation andleadto ©W. WARBY CHAPTER 8 | SYNTHESIS FOR ALL TAXA 267 . et et al cantly dence . (2010) have . 2009). Vollmer . 2009). Vollmer et al c high-confi et al ngerprints of climatengerprints . 2004). cult to produce. The output cult to produce. . (2010) analysis of high-altitude et al et al . 2003; Marshall et al dence for precipitation patterns most dence for precipitation across cation, altering lake productivity (Verburg and Hecky (Verburg lake productivity cation, altering . (2006) analysis of climate impacts on large African Rift impacts on large . (2006) analysis of climate documented similar patterns in Lake Victoria; in some cases,documented similar patterns in Lake Victoria; occurring in deep waters, which shifts are these temperature (Matthews increases air temperature from buffered more are Lake Chad has shrunk signifi recently, 2010). More of multi-decadal declining since the 1950s, in part as a result (Nihoul annual precipitation mean that few impacts from Does the limited literature occurring in African climate change are anthropogenic to infer. This conclusion would be dangerous freshwaters? in Africa is strong, While the paleolimnological literature and amount ofthe limited number of monitoring stations ecosystem, and species data pose baseline hydrological, major challenges that may take decades to correct. and responses trends 8.6.5 Projecting of climate The 21st century should witness the resurgence important change-agent for African change as a more in Africa and (IPCC 2007). Trends ecosystems freshwater conditions will that eco-hydrological suggest elsewhere climate history suggests recent continue to shift, but Africa’s may systems that in African freshwater is resilience that there help with the coming changes. Specifi al lakes or the Eggermont Australian, to North American, mountain lakes. Compared impacts literature African freshwater or Eurasian studies, the thin. remains change studies The most convincing African climate rather than ecological impacts. The document hydrological and Victoria physics of climate impacts on Lakes Tanganyika to changes in and precipitation link shifts in air temperature stratifi 2009; O’Reilly climate shifts to changes in water (2002) has linked recent in Lake Malawi, and Hecky temperature driving African freshwater ecosystems, therefore, comes comes therefore, ecosystems, freshwater African driving paleolimnological and measurements direct a mix of from 2003). studies (Verschuren and proxy fi challenges, the Despite these using both freshwaters detectable in African change are The UN Intergovernmental measures. indirect and direct reviewed (IPCC) has recently Panel on Climate Change impacts generally (IPCC 2007) andAfrican climate change studies have (IPCC 2008). Several for African freshwaters as the Hecky or ecotype impacts, such detailed regional 2010). For the rest of Africa, GCM precipitation projections projections of Africa, GCM precipitation 2010). For the rest projections have proven diffi proven have projections models (GCMs) for Africa show low of global circulation levels of confi latitudes of Africa, with the subtropical and of the tropical exception of the north African–Mediterranean and southern all GCMs show declining levels of where African regions, over this century (Le Quesne annual precipitation cation cation of cult, particularly cations are altering cations are . 2009). et al exibility of extant African species cant portion of agricultural land in Africa cant is the observation that many of thecant is the ed as temporary or ephemeral have been both ephemeral have been ed as temporary or ow regime and hydroperiod. Agricultural water demand and hydroperiod. ow regime are remarkable, particularly for endorheic lake fauna. Genetic remarkable, are studies have not yet fully resolvedand paleolimnological of evolutionary (e.g., rapid evolution and roles the relative responses ecological (e.g., refugial) adaptive radiation) and cichlids, to these pressures Victoria’s by taxa, such as Lake 2008; Elmer (Stager and Johnson orders of magnitude smaller today by the standards of as by the standards today smaller of magnitude orders BP (Issar 2003). years as 7,000 recently systems many freshwater we can infer that By extension, now classifi ecological recent relatively and completely dry over permanent highly are species Given that most freshwater timescales. cycle, the shifts in the hydrological sensitive to even minor and fl ecological resilience Also signifi in Africa haveoscillations between wet and dry periods often on decadal or century rapidly, relatively occurred 2000; Issarrather than millennial timescales (Nicholson drying event that extreme 2003), including the most recent parts of Africa about 4,000 years ago large across occurred and Hooghiemstra 2004). (Marchant climate change in Africa 8.6.4 Signs of anthropogenic in Africa haveThe colonial and postcolonial periods modifi and widespread large seen increasingly dominates water use on the continent. Vörösmarty (2005)dominates water use on the continent. Vörösmarty found that a signifi terrestrial and freshwater ecosystems from irrigation, ecosystems from and freshwater terrestrial mining and other types of intensive resource deforestation, hydropower, pumping, agriculture, extraction, groundwater and industrial and urban development, disease control, manufacturing activities. These modifi conditions in many parts of the region, hydrological and altering quality, changing water connectivity, reducing fl variability and low of high hydrologic is located in areas plans exist for additional modifi Ambitious runoff. sector. particularly for the energy resources, of freshwater which is Grand Inga project, For instance, D. R. Congo’s estimated to be able to generate twice as much electricity dam, and other planned large Gorges Three as the Yangtze’s swathes alter large dams have the potential to profoundly of their basins. The rate of economic development and social change in the past century confounds many of the freshwater that do exist, as dramatic shifts in water use and records management, urbanization, and changes in agriculture of human impacts on freshwater the intensity have altered climate ecosystems. Detecting the signal of anthropogenic change among so many drivers is diffi have had little or no given that many African freshwaters well into the 20th century (Le Quesne monitoring until direct in of climate change into the role 2010). Much research 268 CHAPTER 8 | SYNTHESIS FOR ALL TAXA Data from Ethiopia showhow singledrought events can events —and highpoverty/lowdevelopment levels. precipitation andfl characterized byseasonalprecipitation, highvariabilityin evidence foracorrelation between‘diffi requirements are high.Grey andSadoff (2007) provide of competitionbetweenhuman andecosystemwater Given thatclimatevariabilitymay beincreasing, therisks likely tohaveimportantconservationconsequences. Human responses toanthropogenic climate changeare such droughts are more likelytoappearthiscentury. global climateengines,soitisnotunreasonable toinferthat of thesemegadroughts appearcorrelated with pastshiftsin by relatively abrupttransitions(Shanahan (Scholz decade ‘megadroughts’ across majorregions ofAfrica Several recent studieshavedocumentedsevere multi- Interannual precipitation variabilitymaythusincrease. particularly overthelastcentury(Gergis andFowler2009). past 500years,especiallyextreme ENSOevents,and Oscillation), forinstance,hasbeenincreasing overthe variability attributabletotheENSO(ElNino–Southern GCM approaches (e.g.,Shanahan extreme eventsmaynotbewellcaptured usingexisting climate shifts,havehadinconsistentresults, andveryrare such asdroughts andfl of extreme weatherevents(incontrasttomeanclimate), Attempts toestimatechangesinthefrequency andseverity in signifi the 2050s,whichwilllikelyaltereco-hydrological conditions shifts ofatleast10%meanannualrunoff ordischarge by ecological andevolutionaryphenomenamayexperience of Africanfreshwater fi They concludethatregions nowcontainingmore than80% projected toexperiencelarger shiftsinannualprecipitation. hydrological modellingapproach toidentifyregions (Freeman 2010).Thieme aquifer) watertoAfricanecosystemsandeconomies the importanceofgreen (atmospheric)overblue(surface- 1,000mm inannualprecipitation. Thesesensitivities reveal of 50%forregions thatnowreceive between400and on muchofAfrica,resulting indrops insurfacedrainage annual rainfall(around 10%)couldhaveinordinate effects Stankiewicz (2006)suggestthatevenmodestdeclinesin of freshwater systemsacross thecontinent.DeWitand scenarios forlarge-scale shiftsinmeanannualconditions Several recent studieshaveattemptedtomodeldifferent the future (IPCC2008;LeQuesne especially forinfrastructure lastingmultipledecades into resource managementandinfrastructure designdecisions, of confi know howtointerpret theseprojections andwhatlevel and annualresolutions (Wilby are variableandinconsistentatintra-annual/seasonal dence toplaceinthemasthebasisforguiding cant ways. cant et al . 2007), with some of these episodes marked . 2007),withsomeoftheseepisodesmarked ows, and/orfrequent extreme weather sh diversityandseveraloutstanding oods, asaresult ofanthropogenic et al et al . (2010)usedaclimate- et al . 2009).Itisdiffi et al . 2010). . 2010). cult hydrologies’ — et al . 2008). Climate . 2008).Climate . 2008).Many cult to to cult ‘stationarity’ exhibitedby infrastructure andwater managers Several authors havesuggestedthattheassumptions of that mightenablepositiveecological change? human-induced change,are there actionswecanpursue system iscommittedtodecades orcenturiesofadditional emerging climateconditions.Given that theglobalclimate management regimes fordecadesthatdonotmatch rapid infrastructure developmentmaylockinfreshwater species. TheriskforAfricanfreshwater resources isthat could hastenratherthanbuffer negativeimpactsonAfrican water resource managementapproaches inwaysthat change impactsare likelytointeractwithexistingandfuture millennia. However, emerging anthropogenic climate successfully faceddramaticswingsinclimaterecent Africa’s freshwater ecosystemsandspecies have 8.6.6 et al these riversare shared bythree or more countries(Sadoff Africa hasatleastonetransnationalriver, andabouthalfof boundaries(UNEP2005),everycountryin international regions ofthecontinent.Some64%Africa’s basinscross environmental andresource are governance weakinmany of infrastructure. Bothnationalandtransboundary These challengesare notlimitedtoindividualpieces challenge forAfricanpolicymakers. Le Quesne ‘maladaptations’ toshiftingclimateconditions(Poff 2009; increase speciesvulnerability, andultimatelyleadtogrowing infrastructure mismatchesthatdegradeecosystems, termed ‘stationarity’(Milly conditions; theassumptionofafi design rarely considerclimate-drivenshiftsinhydrological methods of‘sustainable’waterinfrastructure operationsand part ofAfrica’s near-term future. Unfortunately, evencurrent that waterinfrastructure developmentwillbeanimportant (about 30%)relative topotential.Thesefi low levelofhydropower (approximately 5%)andirrigation sanitation needs,butthere isalsogrowing focusonthe Much investmenttodatehasfocusedonwatersupplyand compared to5,961m³intheUSA(Grey andSadoff 2007). quite low—28m³inEthiopiaand687m³SouthAfrica, variability, butpercapitaAfricanstoragelevelsare generally Africa. Water storageisawidespread response toclimate relative topotentialthatexistsacross avarietyofsectorsin noted thelowlevelofwaterinfrastructure development for developmentaid.Forinstance,manyobservershave water managementinAfricaislikelytobeamajortarget development incomplex,oftenunpredictable ways, and As aresult, climatechangewillbeinteractingwitheconomic the endofevent(Sadoff andMueller2009). signifi . 2003). . 2003). cantly lowernationalGDPformanyyearsfollowing freshwater change Implications forfacilitatinghealthyresponses to et al . 2010).Avoiding theseriskswillbeamajor et al . 2008)—canleadtoclimate- xed climate—usually gures suggest suggest gures CHAPTER 8 | SYNTHESIS FOR ALL TAXA 269 exible cant shift in strategy ow regime in lotic systems and in lotic systems ow regime cation to the disturbance regime that the system is that the regime to the disturbance cation cant resilience exists. With thoughtful development cant resilience . (2005) describe a set of strategies to enable climate. (2005) describe a set for many existing conservation plans. for many existing conservation assumes that of an adaptation plan The presence exists for implementation. Conway institutional capacity et al livelihoods.institution-based coping options for East African and information They emphasize better forecasting of fl distribution (e.g., for ENSO), the development managing that can address management regimes resource by events, and preparation for variability and extreme policymakers and governance institutions for long-term climate change. Their suggestions emphasize impacts from to environmental the importance of institutional resilience change. species are The challenges facing African freshwater climate history suggests that daunting, but the continent’s signifi can see a new and management, African freshwaters but not unhealthy qualities. century with different ecosystems, such as high levels of agricultural runoff; the runoff; levels of agricultural as high such ecosystems, that impede as dams such connectivity, of ecological degree of a lotic system; the degree through species movement modifi adapted to (fl ecologically or absence and the presence in lentic systems); hydroperiod are (i.e., climate refugia complex landscape of a climatically of weather events). Evaluation extreme available to buffer assessment a climate vulnerability these elements through a clear climate adaptation plan. The can provide process of these qualities among African and restoration protection would mark a signifi managers resource uencing ned four aspects of ned eco-hydrological . (2010) defi © JEAN-PAUL GHOGUE © JEAN-PAUL et al oor of Lake Chad, the Bodelo Depression is slowly being transformed into a desert landscape. xed or rigidly defi are often widely shared by conservationists as well. Climate often widely shared are on facilitatingchange adaptation, in contrast, must focus species andhealthy change rather than trying to manage ecosystems for fi conditions (Matthews and Wickel 2009; Poff 2009). conditions (Matthews and Wickel 2009; Poff to managing What might a climate-informed approach look like? A full review resources African freshwater but a new vision of is beyond the scope of this chapter, researchers sustainability has been developing among some to climate adaptation and and policymakers. One approach on ecosystemclimate-informed sustainability is to focus past climate qualities that have enabled adaptation through shifts. Le Quesne the level ofecosystems that assist climate adaptation: infl that are detrimental non-climate pressures © JACQUES DESCLOITRES, MODIS RAPID RESPONSE TEAM, NASA/GSFC Once forming the fl Climate driven shifts in hydrological conditions areClimate driven shifts in rarely considered infrastructure when planning large-scale developments that are last many decades. designed to Edea Dam in Cameroon. Chapter 9. Conservation of freshwater ecosystems

Linke, S.1 and Hermoso, V.1 (Section 9.1); Swartz, E.2 and Tweddle, D.2 (Section 9.2); Linke, S.1, Hermoso, V.1, Turak, E.3 and Nel, J.4 (Sections 9.3 and 9.4)¹

¹ Australian Rivers Institute, Grifﬁ th University, Nathan, 4111 Queensland, Australia 2 South African Institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown 6140, South Africa 3 NSW Department of Environment & Climate Change, Sydney, New South Wales, Australia 4 Council for Scientiﬁ c and Industrial Research, Stellenbosch 7600, South Africa

This image of Egypt, taken from the International Space Station, emphasises the importance of freshwater for human civilization. In the view, we see the population of the country almost completely concentrated along the Nile River Valley. © NASA; NASA EARTH OBSERVATORY CONTENTS 9.1 Systematic conservation planning 272 9.1.1 Background 272 9.1.2 Application to freshwater ecosystems 272 9.1.3 Adapting terrestrial approaches to freshwater ecosystems 274 9.1.3.1 Recent developments 274 9.1.3.2 General issues of connectivity 275 9.1.3.3 Threats and lateral connectivity 275 9.2 Site scale conservation 277 9.2.1 Catchment versus site scale approaches 277 9.2.1.1 When are site-scale actions or planning appropriate? 278 9.2.1.2 Prioritisation of site-scale conservation actions 278 9.2.2 Case studies of site-scale conservation intervention in southern Africa 278 9.2.3 Conclusions 282 9.3 Catchment scale conservation 282 9.3.1 Use of biodiversity surrogates 283 9.3.2 Data requirements 284 9.3.3 Planning and implementation spatial scales 284 9.3.4 Freshwater ecoregions as units for conservation planning 286 9.4 Conclusions 286 272 CHAPTER 9 | FRESHWATER ECOSYSTEMS set of areas that cost-effi (Margules andPressey 2000)aimstoidentify anoptimum the mostwidely used.Systematicconservation planning last twodecades–‘systematic conservationplanning’is methodologies havebeendeveloped andappliedoverthe previous conservation planningapproaches, alternative To address thelackofrepresentation andeffi biodiversity needingprotection (Williams However, theyfailtoensure adequaterepresentation ofall et al competes withotherhumaninterests andactivities(Margules not aneasytask.Conservationofbiodiversityvaluesusually Margules threats totheirbiodiversity(Margules andPressey 2000; populations, andbymitigatingatleastsomeoftheproximate contain through maintainingnaturalprocesses andviable should promote thelongtermsurvivalofbiodiversity they varying degrees ofprotection. Onceestablished,reserves common strategyistodeclare theseareas asreserves, with for themaintenanceofecologicalprocesses). The most threatened orrare species,habitatrefugia, orareas important important biodiversityvalues(forexample,areas containing particular areas thatsupport,orhavethecapacityto situation requires identifi freshwater realm heavilyimpacted(Dudgeon of environments (Pimm many different taxonomicgroups andacross awiderange rates are 100to1,000timeshigherthanpre-human levelsin current biodiversitycrisis(Olson The diversityoflifeonEarthisrapidlydecliningunderthe 9.1.1 Background 9.1 Systematicconservationplanning richness orrarity(Williams areas ofhighconservationvalueusingcriteriasuchas Knight 1999;Pressey often havelittleurgent needforprotection (Pressey 1994; efforts onthoseareas thatare easytoprotect, butthat hoc criteria extraneoustobiodiversityconservation.Such on subjectivejudgmentsofbiodiversityvalue,orother et al. potential orsimplyavailability(Margules as scenicvalue,remoteness, lowagriculturalproduction siting conservationareas basedonselectioncriteriasuch conservation wasattemptedby not alwaysbeenthecase.Only20yearsago,biodiversity representative surrogates, onemust remember thatthishas is targeted towards conservingbiodiversityfeatures or While itisanacceptedfactthatconservationplanning biodiversity, isakeystep. of areas intermsoftheirbiodiversity, orcontributionstothat economic perspective.Giventheseconstraints,prioritization to thepersistenceofthosevaluesunfeasiblefrom asocio- conservationstrategiestendtofocus . 2002),oftenmakingprotection ofareas contributing 1996;Sarkar1999).Thesemethodsare basedeither et al . 2002).However, protection ofbiodiversityis et al. cation and focussed management of cation andfocussedmanagementof et al 2000).Othermethodsidentify ciently represent thedesired et al. . 1995), with species in the . 1995),withspeciesinthe 1996;Myers et al ad hoc . 2002). Extinction . 2002).Extinction et al. et al. et al decisionsfor 1988;Pressey 1996). et al. . 2006).This ciency of of ciency 2000). ad threats than adjacent terrestrial ecosystems (Malmqvist Flecker 1993),and are submittedto higher pressures and high amountof theEarth´sglobalbiodiversity (Allanand Freshwater ecosystems containadisproportionately 9.1.2 Applicationtofreshwater ecosystems species whileminimisingthearea oflandrequired. it is,forexample,possibletooptimizerepresentation of chosen oralready protected areas. Followingthisapproach, subsequent planningstepwouldcomplementpreviously and recalculating priorities,sotheareas selected ineach from thedatasetthoseareas thatwere already prioritised not assignedhighpriority. Hesolvedthisbyremoving same time,areas withfew, yetimportant,specieswere for individualspecieswouldbespatiallyduplicated.Atthe recommended forprioritisation,thenconservationefforts (Kirkpatrick 1983;Pressey 2002).Ifalloftheseareas were the highscoringareas were hometothesamespecies reserve coverageofplantspecies,noticedthatmany a Tasmanian botanist,who,whenprioritisingtofi algorithms are basedontheinitialworkofJamieKirkpatrick, performed withcomplementarity-basedalgorithms.These The laststep—maximisingoutcomesiscommonly by Margules andPressey (2000): systematic conservationplanningframeworkdeveloped (1998), andPossingham following keystepsintheframeworksdevelopedbyShea is bestaddressed inadecision-theoretic framework.The and hastheleastimpactonotherstakeholders.Effi social constraints.Thebestplanisonethatcoststheleast adequacy andrepresentativeness underfi comprehensively, whileselectingareas thatensure (CAR). Anoptimalconservationplanprotects biodiversity comprehensiveness, adequacyandrepresentativeness systematic conservationplansaddress three keyprinciples: Although notalwaysexplicitlyacknowledged,most (Margules ad hoc biodiversity features andmore cost-effi complementarity leadtomore effective representations of areas (Possingham of biodiversitywhenasiteisaddedtoanexistingset Complementarity isdefi approaches andincorporatingcostintheselectionprocess. conservation features, usingcomplementarity-based ● ● ●

biodiversity features undertheseconstraints. stakeholders), andtrytomaximiserepresentation of Set constraints(maximumcostorimpacton impacts onstakeholders. List themanagementoptions,theircosts,and This isacrucialstepthatoftenneglected. clear objectivesaboutdesired conservationoutcomes. After collectingdataonbiodiversityattributes,state (Pressey andTully 1994),scoring,orrankingstrategies et al. 2002;Pressey andNicholls1989). et al ned asthegaininrepresentation etal. . 2000). Methods that incorporate . 2000).Methodsthatincorporate (2001)formthecentre ofthe cient solutions than than solutions cient nancial andoften ll gapsin ciency CHAPTER 9 | FRESHWATER ECOSYSTEMS 273

. et al et al. 1991; 2008; . 2004) or et al. et al. et al 2007a; Moilanen sh taxa and assesses relevance sh taxa and assesses relevance et al. 2002). 2011). et al. 2007; Linke et al. es regionally rare ﬁ rare es regionally . 2004), measures of rarity (Filipe . 2004), measures et al. When prioritising for freshwater conservation value, When prioritising for freshwater single species/ towards either geared most studies are Most of or use index-based measures. taxonomic groups richness metrics (Clavero are the index-based measures et al Maitland 1985; Nilsson and Gotmark 1992; Skelton Maitland 1985; Nilsson and Gotmark 1992; Saunders compound richness/rarity/diversity indices (Williams 1995), with a few notable exceptions (Bayley 2004). A more sophisticated approach used an ‘Index of approach sophisticated 2004). A more (Angermeier and Winston 1997), which of Density’ Centres identiﬁ In the last for potential recolonisation. of sites as sources number of conservation an increasing few years, however, starting to are realm in the freshwater planning projects adopt the principles of systematic conservation planning (Nel Hermoso conservation planning might be, in This gap in freshwater systems pose part, due to the complexity that freshwater to conservation practitioners. The connected and complex clear constraints when planning of rivers imposes nature This, combined for conservation in riverine environments.

et al. ne reserve 2008). Most . 2002). Rivers et al et al. 1991; Freitag et al. . 2007), yet they have received . 2007), yet they have received . 2007). Indeed, one of the primary . 2008; Klein et al et al et al 2003). Systematic conservation planning et al. uses of rivers in terrestrial reserves is to defi reserves uses of rivers in terrestrial are 9.1). As a consequence, reserves boundaries (Figure (Lake 1980; features terrestrial towards usually targeted have generally been inadequately dealt with in most ecosystems unless they were assessments of terrestrial biodiversity patterns for terrestrial important considered (Nel and processes conservation planning approaches have, however, have, however, conservation planning approaches because incorporating biodiversity, overlooked freshwater and habitats adds several layers of species freshwater complicated task (Abell 2002). complexity to an already little emphasis on declaring protected has been There purpose of conserving freshwater for the primary areas ecosystems and biodiversity (Saunders methods have been extensively applied to conservation environments and terrestrial in marine problems (e.g., Carwardine and Rundle 2002; Nel the conservation community (Abell little attention from has thrived 2002). While assessment of river condition has argue, in the last two decades and, some would river conservation science efforts, superseded terrestrial and marine assessments is still lagging behind terrestrial of conservation value (Daniels 1997; Root Figure 9.1. Niasse Game Reserve, in the Rovuma River catchment, Mozambique, demonstrating the use of river use of river the demonstrating Mozambique, catchment, Rovuma River in the Reserve, Game 9.1. Niasse Figure river or the catchment of the upper no protection with area, the protected for markers as boundary channels channel itself. 274 CHAPTER 9 | FRESHWATER ECOSYSTEMS upstream ordownstream. Therefore, inaccordance with by activitieshundreds oreventhousandsofkilometres in freshwater systems.Sectionsofarivercanbeaffected 2003), thenature andscaleofconnectivityare different metapopulations (Cabeza2003;FischerandChurch addressing issuesofconnectivitywhendealing with While terrestrial conservationplanningisincreasingly are connectedsystems,bothlaterallyandlongitudinally. confi and riverineconservationplanningistherequired spatial The mostfundamentaldifference betweenterrestrial 9.1.3.1 Recentdevelopments 9.1.3 are notyetfullyappreciated. is alsoclearthatmanyofthevaluesfreshwater species effective conservationpracticesinthefreshwater realm. It impeded ordelayedtheplanningandimplementationof functions, are someofthedrawbacksthatcould have ecological processes required tomaintainecosystem distributions andstatusoffreshwater species,and the with alackofaccessibleandreliable knowledgeon the from space. Congo Riverexits thepoolthroughamarkedlynarrowed channelataseriesofwhitewater rapidsthatcanbeseen in thisview ofthepool(approximately 30kmindiameter).The followsthe southshore boundary known asStanley Pool.Theinternational side oftheriver, endofanalmostcircularwidening inthe withKinshasaonthesouthside.Thecitieslieatdownstream A satelliteimageoftheCongoRiver, demonstratingthecomplexityofriver networks.ThesmallercityisBrazzavilleontheno guration ofpotentialprotected areas. Rivernetworks ecosystems Adapting terrestrial approaches tofreshwater © NASA/COURTESY OF NASAIMAGES.ORG by afewstudiesthathighlightedtheuseoffreshwater river typesandprocesses (Roux — wasaSouthAfricanstudyfocussedonconservingboth a complementarityalgorithmtoderiveaneffi features ofasystematicapproach —clearobjectivesand The fi classifi since Abell dealt withthetopicinboththeoretical andappliedstudies However, inthelastdecadeahandfulofauthorshave freshwater realm. that impededtheearlieruptakeofsystematictoolsin conservation planningframeworkasoneofthekeyfactors of boththeoryandtoolstoresolve thisinasystematic Pereira andCowx2004).We considertheabsence is toodiffi very large streams becausewholecatchmentprotection authors acknowledgethatexceptionshavetobemadefor Crivelli 2002;Collares-Pereira andCowx2004).Some areas (MoyleandSato1991;PuthWilson 2001; conservation valueordevelopingasystemofprotected upstream areas mustbeconsidered whenestimating Hynes’ (1975)paradigmthat‘thevalleyrulesthestream’, rst publicationthatshowedthetwodistinctive cation ofNorthAmerica’s freshwater ecosystems. cult toachieve(Saunders et al. (2000) laidthegroundwork withtheir etal. etal. 2002).Itwasfollowed 2002;Collares- cientsolution river rth CHAPTER 9 | FRESHWATER ECOSYSTEMS 275

et et al. ed a (2007) (2008) 2007a; Nel 2007a) are 2007a) are et al. et al. 2011; Rivers- . 2009) to plan 2009) does not et al. . in press), setting . in press), 2007). Turak 2007). Turak cations have only et al et al. et al. et al. et al et al. 2011; Nel 2008) and a decreasing penalty 2008) and a decreasing et al. 2007a; Linke . 2002; Norris et al. 2011). To date, modifi 2011). To et al. et al (2011) and Esselman and Allan (submitted), (2011) and Esselman and Allan (submitted), cant future advance. cant future ed the extent of present condition and future condition and future ed the extent of present 2011). An alternative has been used solution et al. ed for a freshwater setting, an optimal solution ed for a freshwater et al. et al. cations to two generic conservation planning cations to two generic conservation boundary quality indicator in ZONATION ed the 2007), assuming that land use in the surrounding 2007), assuming that land use in the surrounding 2009; Hermoso 2007b). The most promising approaches to date are to date are approaches 2007b). The most promising conservation along river networks by only considering the conservation along river networks by only (Klein boundaries between connected sub-catchments et al. to existing reserves or headwater streams. These focal These streams. headwater or reserves to existing where of the sub-catchment portions represent areas owing to their importance is most restrictive management Sowa Similarly, biodiversity. for freshwater as fully areas select conservation-opportunity set rules to a number of publications systems. Since 2007, connected into connectivity on incorporating longitudinal focussed algorithms. Others modifi complementarity-based algorithm similar to the one used complementarity-based in C-Plan (Linke upstream entire inclusion of the static rules to ensure riverine plan. Shortly after, catchment in a conservation modifi developed: Moilanen tools were software modifi a number of while connectivity, to deal with longitudinal MARXAN (Ball studies have applied Moore by Turak used to disturbances are of upstream whose aggregations detailed discussion in Section drive connectivity (see more 9.1.3.3). tools can While the above examples show that planning be modifi to conserve the has not yet been achieved. Strict rules entire upstream catchments (Linke upstream entire often unrealistic (and sometimes unnecessary), but only often unrealistic selecting adjacent catchments (Klein automatically guarantee adequate protection (Linke automatically guarantee adequate protection al. rules of connectivity for each the inclusion of different species (Moilanen higher up in the catchment for planning units that are (Hermoso tackled longitudinal connectivity – developing conservation planning tools that consider both longitudinal and lateral linkages in the context of planning for rivers and wetlands will be a signifi lateral connectivity and 9.1.3.3 Threats Lateral connectivity has, in contrast with longitudinal planners by freshwater been considered connectivity, conservation plan every freshwater the start: almost from operates on sub-catchments. Conservation planners have quantifi land-use GIS layers (Linke from threats et al. to biodiversity for threats catchments is a good proxy (see also Stein (2011) integrate lateral and longitudinal connectivity by deriving accumulated land use impacts at every point in model the drainage network, and use a multiple regression contribution of to quantitatively account for the relative . . et et ed et al et al 2007). et al. 2005). The 2005). et al. et al. in press a); and (3) in press . 2007). In addition, rst selected ‘focal’ uenced by longitudinal et al. . (2007) used freshwater . (2007) used freshwater et al . 2004). (2007) fi et al (2007) used C-Plan (Pressey (2007) used C-Plan (Pressey et al et al. et al. . 2007a,b); (2) large-scale ecosystem . 2007a,b); (2) large-scale cation in systematic frameworks frameworks in systematic cation et al sh species in New Zealand (Moilanen sh species in New . 2002; Ward . 2002; Ward et al cation of rivers, whilst Turak and Koop (2008) used and Koop (2008) of rivers, whilst Turak cation 2007), and Abellán . 2008) to identify priority areas for the conservation of . 2008) to identify priority areas sh species to inform input variables in the environmental variables in the environmental sh species to inform input and abiotic data to develop a procedure sh, invertebrate three complementarity-based algorithms were modifi complementarity-based algorithms were three planning for: (1) for use in river systems in conservation taxa in the Australian state of 400 macroinvertebrate (Linke Victoria in all of Australia (Klein processes 18 migratory fi (2005) and Thieme al Addressing aquatic Coleoptera on the Iberian Peninsula. river ecosystems, an example appeared the bias toward that can be suggesting conservation planning approaches applied in to wetlands (Ausseil 2008). 9.1.3.2 General issues of connectivity between planning in the The most obvious difference is, as mentioned above, realms and freshwater terrestrial of rivers and wetlands (Pringle the connected nature conservation 2001). It is nonsensical to design freshwater and downstream without considering the upstream areas as well as catchment ecosystems and land use. areas, The success of conservation actions in any part of a infl river catchment will be greatly connectivity within the catchment. Perturbations upstream reserve the boundaries of a freshwater of or downstream within it will have clear consequences to the processes the long-term persistence of its and its ability to ensure of dams and deterioration of water Presence biodiversity. just quality due to wastewater disposals in a basin are communities apparently freshwater two examples where can be seriously threatened reserves within protected The consideration of far away. operating by processes and its importance in maintaining natural connectivity, waters, is and biodiversity in fresh ecological processes key for systematic conservation planning in these systems (Fausch Over the last few years, scientists and planners have connectivity issues. Higgins started to address and evaluated the connectedness for protection areas year 2007 became a breakthrough year for systematic year for became a breakthrough year 2007 starting systems, planning in freshwater conservation (Abell, reserves framework for freshwater with a new fi classifi fi multi-attribute typology for for generating a bottom-up levels of endangerment and year, rivers. In the same described for river types in South Africa were protection a basis (Nel as targets using quantitative conservation 2007) and an important study in a very data poor area – the a very data poor area an important study in 2007) and Amazon (Thieme de Dios basin in the Madre rivers using developed for classifying were procedures Two multiple attributes: Snelder al. (Fitzsimons and Robertson 2005; Higgins Robertson and (Fitzsimons ecosystem classifi ecosystem 276 CHAPTER 9 | FRESHWATER ECOSYSTEMS stressor issmall, andupstream protection beyondthis 1997). Hence, the sphere ofinfl be metabolised within10kmorless(Storey andCowley scale explicitly. Forexample,mild nutrientenrichmentwill at specifi management zones(seeSection 9.3fordefi and zoningruleswouldbean approach similartocritical The introduction ofstressor- andtarget-specifi landscape scaleprotection schemesneedtobedevised. cannot beprotected, and,inthisinstance,mixedsiteand signifi an internationally fl increases erosion andsedimentationthreats, andreduces downstream from majoragriculturaldevelopment, which Marshes inAustralia.ThisRamsarlistedwetlandis downstream ecosystems.OneexampleistheMacquarie entire catchmentswhentryingtoprotect signifi conservation efforts, asitisseldomfeasibletoconserve lateral mediationofthreats posesthebiggestchallengeto The interactionbetweenlongitudinalconnectivityand complementarity-based algorithm. threats thatmakesdegradedsiteslessattractiveina (2011) created acost-surfaceoutofaggregated upstream local andupstream disturbances.EsselmanandAllan protection ofkeyhabitats. ofthenaturalfl includingrestoration amixoflandscapeandsite-scalemeasures, Ramsar siterequires NSW, Reserve, MacquarieMarshesNature The Northern ofthis Australia.Aswithmanywetlands,effectiveconservation ow through waterextractionforirrigation.Althoughthisis c processes, butwhichaddresses theproblem of cant wetland,theentire catchment © TOMRAYNER uence ofthisparticular nition) directed c connectivity cant between thezonesseems anobviousresearch priority. adaptation for freshwaters thatincludes connectivity background hasbeensetbyAbell contribute toconservationtargets. After thephilosophical to accommodatetheallocation ofmixedzonesthat the software package MARXANhasbeenmodifi ensures maximumeffi be considered together, asonlysimultaneousconsideration management zones(seeSection9.3fordefi focal areas, criticalmanagementzonesandcatchment a posteriori and allocatedcorridorsconnectingthesefocalareas al. of thefewstudiesapplyingtheseprinciples,Thieme addressed infreshwater planningframeworks. Inone stressor-specifi planning frameworks,theideaofmixedusezonesand including riverineconnectivityintosystematicconservation While there hasbeenalotofscientifi be effective conservationmeasures. riparian restoration orsedimentreduction ponds,would planning unit,evensimpleremediation measures, suchas is theonlythreat onasuiteoftargets inadownstream be targeted stressor remediation efforts. Ifsedimentation radius mightbeunnecessary. Anotherexamplewould (2007)selectforthefocalareas inasystematicway, . Inanidealplanningscenario,freshwater c connectivityruleshasonlypartiallybeen ciency. In themarineenvironment, and ow regime et al. (2007),asimilar c progress in nitions) would nitions) ed et CHAPTER 9 | FRESHWATER ECOSYSTEMS 277

. and et al es an et al. Tilapia Tilapia Sandelia ), bluegill ), bluegill c actions c Micropterus Micropterus shes (many and sh populations. is one of four Barbus erubescens ) and banded tilapia ( ) and banded . 2009). This layer identiﬁ Micropterus punctulatus Micropterus c actions. South African scientists et al Pseudobarbus, Galaxias sh sanctuaries (sites or populations) for (Micropterus punctulatus) (Micropterus sh species. c plans can outperform broader regional plans regional c plans can outperform broader ), already present in the lower catchment, the lower catchment, in present ), already ), spotted bass ( ), spotted Lepomis macrochirus (

sh species, such as smallmouth bass ( bass as smallmouth such sh species, sh shes in the National Freshwater Ecosystem Priority Areas Ecosystem Priority Areas shes in the National Freshwater species across the CFR). If the results of Lawler CFR). If the results the species across conservation planning (2003) hold true for most regions, strategies for aquatic ecosystems should include single species and site-specifi have been working on a biodiversity layer for freshwater fi (Nel project (NFEPA) upper limit of 10 fi species and Vulnerable Endangered Critically Endangered, Site-specifi features. biodiversity as irreplaceable in, for example, can also achieve conservation targets catchments, often in conjunction with weakly protected other uses of the catchment. Smaller planning units planning units, and, similarly, usually outperform large site-specifi that might not have clear local goals. alien fi alien dolomieu sunfi sparrmanii where tributary streams and invade to move upstream of populations of smaller cause the extinction they may fi indigenous several undescribed The spotted bass that has alien and invasive bass species in South Africa caused widespread extinction of indigenous fi © SAIAB/ERNST SWARTZ species should for freshwater conservation planning Ideally, using keystone cover whole catchments (see Section 9.3), functioning and species to learn about ecosystem more actions at for monitoring the success of management many species however, the catchment scale. In reality, sub- to single sites or have few remaining restricted are conservation populations – in these cases single species also essential. In the Cape Floristic are programmes 79% of the Region (CFR) of South Africa, for example, fi 43 genetic lineages of primary freshwater with extinction (Linder threatened undescribed) are to only one or 2010). Several species have been reduced two sub-populations or sites (e.g.,

) ) sh may

) Galaxias ) and other Labeobarbus (

ts to other non- sh Barbus serra Barbus erubescens 2003). In the Olifants Austroglanis barnardi c needs of individual sh ( n ( sh ( et al. t an undescribed lled. In some cases, management Clanwilliam yellowﬁ t the Endangered Olifants River lineage Endangered t the ts of goods and services from the services from ts of goods and Pseudobarbus phlegethon

), n ( . 2007) will benefi et al ), and Clanwilliam sawfi t from removing barriers to migration, such as weirs, removing t from ery redfi Labeo seeberi capensis benefi the temporary desiccation of lower stopping and through of water for due to over-abstraction of tributaries reaches allow irrigation. Restoring these corridors will, however, River system on the west coast of South Africa, for River system on the west coast of South for the Critically example, conservation actions planned River redfi Twee Endangered The link between conserving one species. threatened species and the subsequent benefi target not species or ecosystem functions is, however, target must be taken 1998), and care always clear (Simberloff impacts on other species. For to account for indirect example, it is possible that well-intentioned interventions to conserve one species could inadvertently harm 1998). In the Olifants River system, another (Simberloff cyprinids such as Clanwilliam sandfi for example, large ( 9.2.1 Catchment versus site scale approaches versus site scale 9.2.1 Catchment scale, as described planning at the catchment Conservation are, in most cases. There be the ideal approach above, may approach, a catchment scale situations where however, the maintenance of ecosystem aimed primarily at ensuring the specifi functions, may overlook 9.2 Site scale conservation scale 9.2 Site that conservation of ecosystem species. The principle the component and functions will ensure processes healthy (Simberloff also remain species and populations 1998) is not always fulfi ecosystem goods and services may aimed at maintaining but evolutionary history and appear to be successful, a loss through substantially reduced adaptive potential is and future Given the level of current of component species. impacts such as climate species from to freshwater threats (see Section change (see Section 8.6), invasive species it is 8.2.3) and human induced ecosystem degradation, to in order now critical that we conserve species diversity maximise options for the future. of component Maintaining the evolutionary potential in the more species while society is primarily interested immediate benefi is perhaps the biggest challenge facing global environment ‘less important’ Apparently conservation programmes. intrinsic rights species should be valued not only for their opportunity values as to exist, but also for their future Such species may also act important genetic resources. for the conservation of many other surrogates as effective Lawler species (e.g., non-target (Impson species with a similar distribution and conservation status. Prioritising conservation of populations of the Critically catfi rock Barnard’s Endangered will similarly benefi of fi 278 CHAPTER 9 | FRESHWATER ECOSYSTEMS to understand evolutionary patterns andprocesses.to understand evolutionary patterns species distributions, andstudiesofgenetic diversity it requires fi however, bediffi information isavailable.Obtaining thatinformationcan, plan canberelatively simpleoncethenecessary incorporation aspartofawider catchmentmanagement Choosing sitesforsite-specifi actions forthreatened speciesshouldbethepriority. extensive intervention,thesimplerandcheaperconservation where capacityandresources are notavailableformore site-specifi features orprocesses; and2)through prioritisationbetween to theextinctionorunacceptablelossoflocalbiodiversity in twosteps:1)where afailure tomeetthe target willlead Site-specifi 9.2.1.2 Prioritisationofsite-scaleconservationactions expensive interventionsmaybetheonlyoption. result insignifi cases, smallchangesinlanduseorwatermanagementcan site-specifi catchment-wide planning(prioritisationofecosystems)with their survival. species maynotalwaysbeprotected intheoptimalsitesfor planning isonacatchmentscale,highprioritythreatened may notbewithinthearea ofoverlap.Therefore, where protect specifi although theseareas are notnecessarilythebestplacesto highlighting areas ofoverlapbetweenthreatened taxa, Catchment-wide planninghasthefurtheradvantageof effective (forexample,becauseofmore extantpopulations). threatened speciesforwhichconservationactionmaybemore biodiversity areas maybeneglected,aslowerpriority The disadvantageofsite-specifi and willoftenbenefi effectively, are achievableinareas where capacityislow, for localscaleactions.Suchtargets canoftenbemetcost understandable andeasilyobtainabletargets canbeset on-site. Theadvantageofsite-specifi evolutionary history. Solutionsmust,therefore, befound indirect threats tootherbiodiversity, andloss ofaregion’s 1997), translocationofparasites(Kennedy1993),direct or 1992; Leary genetic integrityofpopulations(DowlingandChilds translocation, canitselfcreate problems, suchasaffecting no optionsforprotection elsewhere. Theotheroption, of aspeciesorecosystemiscritical,there are usually few sitesthatcanbeclearlydefi IUCN CategoriesandCriteria)orrestricted tooneor very Critically Endangered orEndangered, according tothe are highlythreatened (usuallythosespeciesassessedas Site-specifi 9.2.1.1 appropriate? When are site-scaleactionsorplanning c actions withinoverallcatchmentplans.Inareas c actions(criticalspeciesconservation).Inmany c conservationactionsshouldbeprioritised cplanningandactionsare neededwhenspecies eld survey, knowledge ofknownandprojected et al cant conservationsuccess,butinothercases

c taxabecausetheirhealthiestpopulations An overallstrategyshould,therefore, combine cult, time consuming andexpensive,since . 1993;Quattro t otherspecies. c interventionsisthathigh et al ned. Whenthestatus c interventionand c interventions isthat . 1996;Avise et al . without successful recruitment. Clanwilliam yellowfi smallmouth bass species, catfi ( for example,theindigenousfi alien smallmouthbassisfound. IntheRondegatRiver(), species survivingbelowthebarrierwhere theinvasive found aboveasmallnaturalbarrier, withnoindigenous In atypicalscenario,fi indigenous fi eradication ofinvasivealienfi within abroader ecosystemconservationframework,where a goodexampleofsite-specifi Conservation andSustainableDevelopmentproject provides The CapeActionPeopleEnvironment (CAPE)Biodiversity of SouthAfrica fi Development Project:Eradicationofinvasivealien 1. C.A.P.E. andSustainable BiodiversityConservation 9.2.2 benefi from themostthreatened toleastthreatened andacost- to speciesconservationstatus.Speciesshouldberanked have beenidentifi when prioritisingspeciesforconservation.Oncesites Evolutionary uniquenessshouldbeamajorconsideration alien fi CFR havesomeofthemostextreme examplesofinvasive Cederberg tributariesoftheOlifantsRiversystemin highlighted ashavingamajorimpactinalmostallcases. alien fi (http://www.iucnredlist.org; Tweddle this region are threatened, according totheIUCNRedList © SAIAB/ERNSTSWARTZ upstream. smallmouth bassfromspreading The RondegatWaterfall, theinvasive whichprevents Barbus calidus shes fromriversectionsintheCapeFloristicRegion s ( sh t analysisconductedtoprioritiseactions. intervention in southern Africa intervention insouthern Case studiesofsite-scaleconservation shes introduced from thelate1800stomid1900s shes impactingpopulationsofindigenousfi

Austroglanis gilli survive aboveasmallwaterfall thatprevents sh faunaoftheCFR.Almostallﬁ ), Clanwilliamyellowﬁ

from spreading upstream. Onlyafewlarge ed, actionscanbeprioritisedaccording sh

are able tosurvivewiththebass,but ve tosixindigenousspeciesare )

and anundescribed ery redfi ery shes willgreatly benefi c conservationtargets set et al sh, Clanwilliamrock n, Clanwilliam redfi . 2009), with invasive . 2009),withinvasive sh speciesin Galaxias Galaxias shes. shes. t the the t n CHAPTER 9 | FRESHWATER ECOSYSTEMS 279 shes once once shes shes in these shes constituted a © SAIAB/ERNST SWARTZ ed four river sections for a sh. rst because of the ease of treatment rst because of the ease of treatment shes from the Clanwilliam Reservoir on shes from shes was feasible; (c) the rivers would to the indigenous species; (b) eradication cant threat process, involving all conservation scientists with process, knowledge of the CFR, identifi pilot project where: (a) alien invasive fi where: pilot project signifi of invasive alien fi healthy habitats for the indigenous fi still provide and (d) the rivers were removed; the invasive aliens were alien targeting not of major importance for angling, often invasive species. the invasive alien fi to eradicate A proposal as the main river sections, with the piscicide rotenone Environmental eradication method, underwent a rigorous 2009). This recommended Impact Assessment (EFA ahead, with very strict environmental go that the project that the Rondegat River recommended The EIA safeguards. fi should be treated with the lessons and the minimal risks to the environment, The learned eradication programmes. applied in future of the EIA have been accepted by all recommendations being is authorities, and the project environmental relevant implemented. on the Rondogat River involves The pilot project upstream farm weir that prevents an existing strengthening migration of alien fi the main stem of the Olifants River immediately downstream 9.2). Eradication of river (Figure stretch of the targeted shes) sh populations. sh sh eradication strategy. A selection sh eradication strategy. shes (thick grey line). The area between the line). The area shes (thick grey The need for a programme to protect the indigenous fauna to protect The need for a programme and was subsequently initiated in of the CFR was clear, the form of an alien fi Rooidraai Waterfall and the downstream weir will be and the downstream Rooidraai Waterfall (eradication of alien fi targeted for rehabilitation The stretch the Rondegat River that is to be treated of to eradicate alien species of fi with piscicide 9.2. The Rondegat River and Clanwilliam Dam, Figure indigenous showing the distribution of most of the fi to expand the indigenous fi 280 CHAPTER 9 | FRESHWATER ECOSYSTEMS for thetownofBarrydale.Thespeciessuffers mostof been furtherfragmentedbywaterabstraction,mainly the onlyindigenousfi 2005). Therestricted rangeofthisspecies,whichis in theWestern CapeProvince ofSouthAfrica(Swartz site intheTradou catchmentoftheBreede Riversystem This undescribedBarrydaleredfi South Africa (Pseudobarbus sp.) redfi actionsfortheBarrydale 2. Conservation existing naturalbarrierupstream. Clanwilliam Reservoirbelowtheweirandverysmall species, andabuffer zonebetweenthealiencolonised outcome willbetheextendeddistributionofindigenous species from theircurrent restricted habitatupstream. The recolonisation bythefi and thesmallwaterfallupstream willallowsubsequent of aliensfrom the4kmstretch ofriverbetweentheweir © SAIAB/ERNST SWARTZ (Pseudobarbus sp.) redfi Barrydale The CriticallyEndangered (thick grey line),duetochangesinwatermanagement andalienfi Figure 9.3.TheTradou catchment,showing thechangeindistributionofBarrydaleredfi are being threatened by mainlyalienfi beingthreatened are . Manyotherredfi intheCapeFloristicRegionof sh speciesinthecatchment,has ve indigenousthreatened fi n isrestricted toasingle n specieslikethisone sh species. sh n n sh tributaries restore habitatinthelowerreaches ofitsrange. remain intheupperreaches. Somereturnfl above thenaturalrangeofspecies,andonlypools range. Excessivewaterabstractionstopstheriverfl sparrmanii sunfi Alien largemouth bass( the threats commontoindigenousfi picture) and some upstream areas. areas. and someupstream picture) redfi in thecompleteextirpationofBarrydale species, resulting The Tradou RiverGorgehasbeeninvadedbyalienfi n inthelowersectionsofgorge (asseeninthe sh ( sh sh distributions. Lepomis macrochirus ) haveinvadedmostofthespecies’indigenous Micropterus salmoides ) andbandedtilapia( © SAIAB/ERNST SWARTZ n ( Pseduobarbus shes oftheCFR. ow andsmall ), bluegill sh Tilapia sp.) ow CHAPTER 9 | FRESHWATER ECOSYSTEMS 281 c et cation of c action is sh species, the 2004). , a Critically et al. oodplain in the dry season. 2004). The streams inhabited 2004). The streams (Neolebias lozii) et al. cial barriers to protect it from threats threats it from cial barriers to protect . 2002) been implemented in full, the . 2002) been implemented in Zambia et al ooded plains (also called dambos) bordered ooded plains (also called dambos) bordered n’s Red List status might well have been Red List status n’s oating mats of vegetation adjacent to the stream stream oating mats of vegetation adjacent to the . 2001; Rall The banded Neolebias The banded Neolebias © DENIS TWEDDLE Endangered species from the Kataba River and its tributary, Endangered River and its tributary, species from the Kataba the Sianda, a small tributary system of the Upper Zambezi, which it joins on the eastern the Barotse Floodplain. side of In contrast to the previous examples of site-specifi In contrast to the previous conservation in southern degradation of the Africa, where of human activities has as a result aquatic environment localised interventions, the banded necessitated urgent Neolebias does not need immediate action to reverse degradation, or artifi such as alien species. Instead, site-specifi to the posed of the threats awareness needed to promote unintentional degradation species’ survival, and to prevent is low of the habitat. The human population in the area in the dambos poses the agriculture and but increasing, the from Part of the Kataba River upstream threat. greatest known location of the species has been canalised, and no found in this section of the river, banded Neolebias were removed as the dense mats of peripheral vegetation were also when the canal was dug. Canalisation of the streams the fl from accelerates run-off population leading to intensifi human Increasing In the Upper Zambezi River system, one fi In the Upper Zambezi River system, one This species is banded Neolebias, is Critically Endangered. to the Kataba River and its tributary, restricted apparently as intensive sampling in apparently the Sianda stream, of the Upper Zambezi the rest suitable habitats throughout These two system has failed to yield any other populations. form a very small tributary system of the Upper streams Zambezi, which joins on the eastern side of the Barotse Floodplain (Tweddle small and associated with by the banded Neolebias are seasonally fl in and under by open woodland. The species occurs dense fl vegetation (Winemiller and and in dense emergent margin Kelso-Winemiller 1993; Tweddle al Maloti redfi due to the down-graded, but this is not yet possible, threats. remaining 4. Conservationfor Banded Neolebias plans (Neolebias lozii) species. Had the research recommendations (Skelton recommendations species. Had the research sh cial n may species rst canal n could be n could . 2002). Two . 2002). Two sh predators predators sh et al (Pseudobarbus n n. Water is currently is currently n. Water Pseudobarbus . 2002). Recognising that the unpublished). Predatory fi unpublished). Predatory et al et al. n is a Critically Endangered species species Endangered n is a Critically in Lesotho . 2001; Rall . 2001) that has evolved without fi ns themselves could pose a threat to the other could pose a threat ns themselves et al et al ows required under South African legislation. ows required . 2001). As a mitigation measure, a dual approach a dual approach . 2001). As a mitigation measure, n. The Mohale population is genetically different to n. The Mohale population is genetically different n, and, in addition, it is possible that the redfi (Skelton the highlands of in the headwaters of the Orange River in the system as Lesotho. Mohale Dam has been built within to Gauteng part of a water transfer scheme to deliver water in South Africa. The dam was unfortunately built Province to the future on an important catchment with regards extant population of the Maloti survival of the largest redfi distance similar the other four populations, with a genetic in other to species level differences (Swartz 2005; Swartz The Maloti redfi In this case, simple interventions could greatly improve improve greatly could interventions case, simple In this redfi the Barrydale for the situation a canal and diverted through upstream from removed town and farms (Figure delivers water to the system that back into the river is released for the farmers 9.3). Water down. If into canals lower it is again diverted where from bypass the fi to farmers could the water allocated lower downstream, in the river until abstraction and remain for the Barrydale redfi at least 1km of habitat the upper reaches bass occur in Largemouth restored. the disappear from will probably of the catchment, but dams in the upper catchment. There from river if removed local community in also opportunities to involve the are formerly played. children where cleaning up their river, a contentious remain will, however, abstraction Water being water in the river is currently issue. Even though all and there of Barrydale is increasing, used, the population The site-based water restrictions. severe already are some respite, above will provide actions recommended of the but a lasting solution is needed to allow restoration river fl 3. Lesotho Highlands Development Authority project to conserve Maloti redfi the quathlambae) the species can now invade the Mohale Dam through water transfer tunnel, with potential impact on the Maloti redfi not be able to adapt to the new lacustrine habitat (Skelton et al that part of the natural range was adopted to: 1) secure of the Mohale population above the dam with an artifi barrier; and 2) translocate individuals to neighbouring alien invasions and habitat alteration from catchments free (Skelton of the four translocations seem to have been successful. the native barrier to protect the proposed Unfortunately, Mohale population was not built, leaving the natural range of this unique lineage still vulnerable to invasion by alien Maloti redfi aquatic fauna of the translocation sites, an invertebrate only minor study was conducted. The study predicted impacts, as no invertebrates of conservation concern, and found (Rall no endemic species, were 282 CHAPTER 9 | FRESHWATER ECOSYSTEMS always be preferable, butwith manyspecies restricted can belinked.Catchment-wide conservationplanning will for aquaticbiodiversity towhichclearconservation actions planning, itis,therefore, importanttoincludecriticalsites of betterinformation(Tweddle changes instatusbutare insteadmainlyduetoprovision been down-graded,althoughthese donotrefl as adirect result ofconservationactions.Somespecieshave removed from athreatened categoryontheIUCNRedList AfricanfreshwaterNo southern fi that isatgreat riskofbeinglostforever. services butmayoftenholdsignifi to provide littlebenefi of site-scaleactionare thosespeciesthatmayappear management actions.Amongmanypotentialbenefi to speciesthatmightotherwisenotgainfrom large scale Simple actionsatthesite-scalecanprovide rapidbenefi for thetargeted conservationofrange-restricted species. framework ofcatchmentscaleinterventions,inparticular Site-scale conservationhasitsplacewithinthebroader 9.2.3 Conclusions canals orclearingtheswampvegetation. to maintainthediversestream habitatandavoiddigging in theirriver. Hopefully, peoplemightthenbeencouraged communities, whomaynotknowthatthisrare specieslives is recommended toraiseawareness amongthelocal the ZambeziRiver, therefore, site-specifi is alsoapotentialthreat. Inthistinysub-catchmentof that anyreduction infl large headwaterswampsustainstheKatabastream, such and herbicides,islikelytothreaten thehabitatfurther. A bordering thedambos,andpossiblepollutionbyfertiliser agriculture, togetherwithdeforestation ofthewoodland banded Neolebias Sampling adensematofvegetationintheKatabaRiverusingrobustD-net.Thisyieldedseveralspecimens (N. lozii) ow duetogroundwater abstraction t through provision ofecosystem . © DENISTWEDDLE et al . 2009).Inconservation cant geneticdiversity shes haveyetbeen c intervention c ect genuine genuine ect ciaries ts interventions. funding are available,havethepotentialtoinspire larger success stories,inparticularwhere limitedcapacityand to haveamajorbenefi Africa,buttheresouthern are opportunitiesforsuchactions protect threatened aquaticspeciesare intheirinfancy irreplaceable features. Site-specifi to onlyoneortwosites,itisessentialincludethemas species (seeDarwall The increasing availabilityofspatialdatasetsforfreshwater fortacklingtheseproblems.alternatives stages. Inthissection,we will discusstheavailable essential forintegrationofplanningandimplementation be carriedout.Selectionofthemostappropriate scaleis the spatialscaleatwhichconservationexercise will Secondly, andnolessimportant,itiscrucialtodefi represent thespatialdistributionofconservation features. of approaches are beingusedinconservation planningto of areas thatrepresent allthetargeted biodiversity. Arange is required toidentifythemostcost-effective combination distribution ofbiodiversityfeatures andconservationcosts planning isusuallyaspatially-explicitexercise, sothespatial spatial distributionofconservationfeatures. Conservation in scientifi on-ground issues.Themostbasic,anddiscussed planning describedabove(Section9.1)—posessome scale —inlinewiththeprinciplesofsystematicconservation Planning andimplementingconservationatthecatchment 9.3 Catchmentscaleconservation freshwater systems. of systematic conservation planningapproaches in here forAfrica,are openingthe door forapplication c literature, istheneedforinformationon et al t forthreatened species.Local . 2010),suchaspresented c conservation actions to cconservationactionsto ne ne CHAPTER 9 | FRESHWATER ECOSYSTEMS 283

et ne et al. et al. 2007) (2011), et al. et al. c defensibility (2002), Higgins . (2011), Nel 2004; Higgins et al. These use abiotic et al ned on the basis of aned on the These are types ofThese are (2011). et al. (2007), Ausseil et al. cations are confounded by cations are cation to within the boundaries cation ts a single environmental model environmental ts a single et al. cations are usually hierarchical and usually hierarchical cations are ned biological attributes, for example,ned biological attributes, for (submitted), Khoury ed physical surrogates has been heatedly ed physical surrogates et al. (2005), Thieme ciency stemming from certainty of protecting a certainty of protecting ciency stemming from 2003). al. (2011) and Rivers-Moore (2011) and Rivers-Moore suggesting that without calibration 2004), with research scale, such by observational surveys at the appropriate be applied with caution (Lombard should surrogates information only and are commonly applied in data- information only and are limited biological with severely poor planning regions deriveddata and expert knowledge. These GIS make use of best available data and surrogates freshwater conceptual knowledge on abiotic drivers of systems. Such classifi to abiotic variables to defi assigning thresholds knowledge about how ecosystem types. This requires evolutionary species, communities, and ecological and to the physical and chemical respond processes limited knowledge Based on the current environment. ecosystems, the scientifi of freshwater is dubious. In addition, the utility of such thresholds of unstratifi (Araujo and marine realms debated in the terrestrial and Williams 2001; Brooks et al. Heiner development, as it fi as development, data. to poor input robust and is species to multiple Dissimilarity is similar to Generalised This technique used in the context (GDM), which has been Modelling (Ferrier conservation planning of terrestrial turnover which models species as a technique along By of actual species. gradients instead environmental the turnover, for responsible identifying the variables that would not otherwise be features even very rare in the conservation plan. considered modelled are defi ecosystems or landscapes and physical attributes. Thesewide range of biological ecosystem classifi constrain abiotic classifi of coarsely-defi that are ecoregions zoogeographic zones or freshwater (but not exclusively) expert-derived. Examples frequently include Roux of such approaches Choice of surrogates ultimately depends on the availability Choice of surrogates scale, as well as the expertise of data at an appropriate for undertaking the assessment. Rapid and resources where assessments or assessments in data poor regions no species or observational data exist will only be able to the – recognising physical surrogates make use of pure (see above) and tempering limitations of this approach with expert knowledge. However, possible it wherever based on actual the key advantage of using surrogates for Africa, is in this report biological data, as presented their effi 4 physical surrogates. Pure 3 Ecoregional surrogates. surrogates. 3 Ecoregional

et This 2008; 2005). et al. c data (Abell et al. . 2010), the technique by 2003; Wilson 2007b; Moilanen et al 2011) based on environmental 2011) based on environmental et al. et al. . 2011) or communities (Turak and Koop and Koop . 2011) or communities (Turak et al. et al ed by relating them directly to biological survey them directly ed by relating ciency (CARE). Therefore, any surrogates used in any surrogates ciency (CARE). Therefore, cation of biodiversity in a planning unit – in this unit – in this biodiversity in a planning cation of (2007) and the study by Leathwick (submitted). category includes species, species assemblage category includes species, species Biological survey data types, or observed processes. usually fraught with data gaps (false absences). are use of inconsistent survey data in conservation Direct areas planning will bias selections to well-surveyed un-surveyed areas and ignore of the planning region, bias this address be important. To in reality, that may, continuous geographic coverage, and obtain a more species many practical applications have modelled distributions (Linke Leathwick (submitted) is a particularly exciting new Hermoso 2008; Turak al. Given the lack of freshwater-specifi 2002; but see Darwall attributes. While this alleviates the problem of false attributes. While this alleviates the problem challenge – that absences, it brings with it a different In conservation planning, false of false presences. in that a problem are errors, or commission presences, they lead to the false assumption that by capturing you have it is falsely present a species a site where conserved (Loiselle These are environmental surrogates that have been that have been surrogates environmental These are stratifi no unlike in the above category, data. However, Instead, predicted. directly biological attributes are surrogates a statistical model links environmental to the landscape patterns of biological attributes. by Snelder provided are examples Freshwater . (2011) suggest the following taxonomy of surrogates: . (2011) suggest the following taxonomy of conservation planning need to relate to the features we want to the features conservation planning need to relate spatial and temporal scales. Linke at the relevant to protect, et al 1 based on biological survey data. Surrogates 9.3.1 Use of biodiversity surrogates surrogates Use of biodiversity 9.3.1 complete as we never have a need to be used, Surrogates quantifi useful (2002) describes a Sarkar case, a sub-catchment. and ‘estimator between ‘true surrogates’ dichotomy species or target the are surrogates True surrogates’. targets hence ideal ‘direct’ themselves and are processes as every researcher However, in a planning framework. it is almost impossible to get a or practitioner knows, all planning across true surrogates complete coverage of situations, we need an life in most real units. Therefore, that – features estimator surrogates informed selection of species or processes. target the true represent need to we of surrogates, evaluate the best selection To should help the surrogates keep in mind the objectives: on principles ofus design a conservation plan based Representativeness Adequacy, Comprehensiveness, and Effi 2 Biologically-informed physical surrogates. 284 CHAPTER 9 | FRESHWATER ECOSYSTEMS extremely limited andthearea very large. Akeyadvantage 2007) orChina(Heiner in extreme cases,forexampleintheAmazon (Thieme the useofrawenvironmental surrogates isonlynecessary modelling techniquesthatare robust todatapoorinputs, approaches. Moreover, withtheavailabilityofpredictive and especiallynoreason tojustifynotusingsystematic planning isnotasmuchofan issueasitusedtobe, data availabilityforsystematicfreshwater conservation informed physicalsurrogates. Hence,weconcludethat easier toproduce modelledtruesurrogates orhighly friendly modellingtechniques,itisrapidlybecoming in thisreport forallofcontinentalAfrica,andnew, user- With increasing availabilityofbothGISdata,asprovided one haslookedelsewhere. only siteinthelandscapewiththisfeature, butbecauseno not beensampled–necessarilybecausethisisthe favour siteswhere dataexistandignore sitesthathave limitations. Forexample,scoringapproaches willalso et al. been acknowledgedinsystematicplanning(seeRondinini argument: whilebiasfrom incompletedatacoveragehas assessments are preferable. However, thisisnotavalid data hungryand,therefore, index-basedbiodiversity systematic planningapproaches isthattheyare too issue ofdataneeds.Afrequently heard criticismof Closely linkedwiththediscussionofsurrogates isthe 9.3.2 Datarequirements can readily beappliedtodata-poorregions (Ferrier2002). beds forevaluatingtheperformanceofsurrogates that it alsomaybepossibletousedatarichregions astest- largely onremotely defi poor regions, defi that is,betaandgammadiversity(Ferrier2002).Indata- particularly usefulforrepresenting differentiation diversity, also beenargued thatcommunity-levelsurrogates maybe account forunknownandunder-surveyed species.Ithas can becombined—isdesirable,becausethismayhelp Guisan 2006)—especiallyifseveralcommunitytypes process. Usingcommunity-levelsurrogates (Ferrierand Africa willprove extremely usefultoinformtheplanning preferable –onceagain,datasuchasprovided here for possible, usingsurrogatesis basedonobservedpatterns to protect onlythehighestprioritysitesso,wherever most places,resources are, however, likelytobeavailable could thenbeselectedatalowlevelofthehierarchy. In resources are unlimitedbecausethefeatures toprotect 2002) mayensure thepersistenceofbiodiversitywhere geology andhydrological boundaries(Snelderand Biggs 2005) oraclassifi (Frissell hierarchical classifi plans usingonlysurrogates defi given amountofbiodiversityperunitarea. Conservation 2006),scoringapproaches ultimatelyfacethesame etal. 1986;Thoms nition ofsurrogates willinevitablyrely cations basedongeomorphology cation thatincorporatesclimate, etal. ned features withlowcertainty, but 2011),where speciesdataare etal. 2004),hydrology (Stein ned bytop-down etal.

highly relevant in the contextof multiple-use freshwater hierarchical protection strategy byAbell the biodiversity itcontains.Inthisrespect, therecent whole catchmentsbutfi at thecatchmentscaledoes notmeanprotecting attention beingdedicatedto this keyissue.Planning at thecatchmentscalehaveprevented more effort and and thelackoftoolstoaddress conservationplanning misconception ofplanningandimplementationscales incorporated intheconservationplan.However, the processes, such asmigrations,canbemore effi (e.g., protect wholecatchments).Thisway, catchment not necessarilyhavetobeimplementedatthatscale the wholecatchmentscale,aswestateabove,butdoes successful, aconservationprogram shouldbeplannedat planning andthescaleofimplementation.To be two conceptsare usuallyconfounded:thescaleof When consideringspatialscalesinplanningprocesses, 22, thisvolume). Lake Victoria (MumbaandHoward 2005)(butseepage consequences onendemicfreshwater fi and Niletilapia( these introductions, suchastheNileperch ( distant areas (Clavero andGarcia-Berthou 2006).Someof between neighbouringcatchmentsorintroduced from example, freshwater speciesare constantlytranslocated been weakenedbyhumaninterventions,andnow, for neighbouring riverbasins.Thisimportantrole hasrecently also limitthedirect transference ofthreats between (Avise 1990;Verheyen endemics thatevolvedfrom onlyafewcommonancestors lakes), withmore than350speciesofcichlids,mostthem the geographicalisolationofcatchments(eitherriversor 2007). LakeVictoria isanexampleoftheimportance as speciation,occurwithincatchmentboundaries(Rahel processes occur. Importantecologicalprocesses, such be considered biogeographicislands,where speciation pose impassablebarrierstothesespecies,hencetheycan and thepropagation ofthreats. Catchmentboundaries low inter-basin connectednessforstrictlyaquaticspecies propagation ofthreats alongriverbasinscontrastswiththe The strong intra-basinconnectivitythatfacilitates the 9.3.3 Planningandimplementationspatialscales time, givenbothnaturalandhumandisturbances? freshwater biodiversitypersistsandevolvesnaturally over adequacy ofconservationplans–howdoweensure that for freshwater conservationplanningliesinaddressing the regions), webelievethatamuchmore signifi in conservationplans(particularlythosedatapoor species andhabitatswillimprove therepresentativeness While continuedinventoryingoftheworld’s freshwater available andgeneratereliable butcoarseassessments. of existingdata.Theycanbeappliedwhere littledata are of systematicapproaches isthattheymakethebest use Oreochromis niloticus Oreochromis et al nding thebestway tomaintain . 2003).Catchmentboundaries sh communitiesin ), havedrastic et al. cant challenge Lates niloticus (2007)is ciently ) CHAPTER 9 | FRESHWATER ECOSYSTEMS 285 2007). et al. Figure 9.4. Schematics of proposed freshwater protected area zones. (a) Freshwater focal areas, such as such focal areas, (a) Freshwater zones. area protected freshwater of proposed 9.4. Schematics Figure or populations, species, focal supporting or wetlands streams, headwater lakes, reaches, river particular habitats or upstream connecting key as river reaches zones, such (b) Critical management communities. (c) A catchment areas. focal freshwater to the function of will be essential whose integrity riparian areas, or focal area freshwater the most downstream of upstream catchment the entire zone, covering management would be applied principles catchment management within which integrated zone, and critical management Abell from with permission (reproduced 286 CHAPTER 9 | FRESHWATER ECOSYSTEMS them. Imagine ifaspecies orotherecosystem attribute increases uncertaintyaround within spatialpatterns within ecoregions. Heterogeneity inlarge planningunits planning frameworks.Akeyproblem isdataheterogeneity used asmeaningfulplanning unitsinconservation high-level surrogates —theycanbetoocoarseto to bothstratifyreporting units—aswellproviding single unit.However, althoughecoregions canbeuseful species assemblagesandecologicalprocesses intoa basin, orforaggregating severalbasinssharingsimilar meaningful unitsforsubdividingalarger riverorlake as conservationplanningunits–biogeographically this volume).Theseecoregions are designed toserve particular (Thieme ecoregions intheWorld (Abell been recently devotedtothedescriptionoffreshwater different biodiversitysurrogates. Substantialefforts have spatially-explicit informationaboutthedistributionof the continentalscale.Theyare usuallydefi and processes across large spatialscales,suchas inspeciesassemblagecomposition,habitats, patterns for therepresentation ofnaturaldistinctiveness Ecoregional classifi 9.3.4 entire upstream zones. mixed-use protection schemesmightnotberealistic for Congo andNileRiverbasinsinAfrica.Inthesesystems, the Murray-Darling(Australia),Mississippi(USA)or very large systems,suchastheAmazon(SouthAmerica), However, more thoughtneedstobedirected towards without beingtoorestrictive across theentire catchment. the termemphasizesawhole-catchmentperspective zones are similartomultiple-useprotected areas, but management principlesapply. Catchmentmanagement groundwatersheds (Pringle2001),basiccatchment area. Inthistrulymixed-usezone,whichalsoprotects the entire catchmentupstream ofthefreshwater focal ‘catchment managementzone third, andfi at thetimesrelevant tospawningandmigration. The environmental waterallocationsmayonlybeneeded spawning migration,aswelltemporal sediment load,keyfi management zonesmaybespatial,suchasareas of high downstream freshwater focalareas. Examplesofcritical managed tomaintainthekeyprocesses thatsupport the designated as‘criticalmanagementzones’,whichwillbe be protected, someofthekeyareas upstream wouldbe If —asinmostcasestheentire catchmentcannot similar toterrestrial protected areas (seeSection9.2). and usewithintheseareas islikelytobefairlyrestrictive, biodiversity wouldbedeclared a‘freshwater focalarea’ high priorityareas fortheconservationoffreshwater zones (Figure 9.4).According conservation planning Freshwater ecoregions asunitsfor nal, leveloftheprotection strategyisthe et al cations provide appropriate units sh spawninghabitatorcorridorsfor . 2007)(seeChapter1,Figure 1.2, ’, et al whichisdesignatedto to thisnewschedule, . 2008)andAfricain . Forexample, nd using ned 4 3 2 1 large planningunitsforanumberofreasons: spatial information.Smallerplanningunitsoutperform here forAfrica—itdoesnotmakesensetoignore fi available —suchasatthesub-basinscale,presented representation. Inthecasewhere more detaileddataare data can,inparticular, leadtomassiveshortfallsofreal ecoregion intothesmallerunits.Patchydistribution it followsthatthispresence istranslatedfrom the If anecoregion isnowsplitintosmallerplanningunits, unit. Usuallythisisreported as‘present’ intheecoregion. ofalargeoccurs justinthecorner ecoregional planning conservation managers, stakeholders andthescientifi the biodiversity containedwithinhasprobably prevented catchments have tobe‘lockedup’inorder topreserve continental Africaare already available.Theideathatwhole for more than7,000sub-catchments across thewholeof current report, dataonspeciesstatusanddistribution have beengathered. Forexample,aspresented inthis on biodiversitydistributionand environmental surrogates planning needs.Inthelastdecade,extensivedatabases the pretext ofalackthedetaileddatathatsystematic protecting biodiversityeffi criteria approaches thatfallshortatrepresenting and the biodiversitytheycontain.Useof planning willenhancethecapacityofreserves tomaintain Incorporating theseecologicalaspectsinconservation the tools(MARXAN,ZONATION) andmethodsavailable. along rivernetworks,canbeeffectively addressed with of threats (forexample,pollutantsorinvasivespecies) ecosystems, suchasmigrationorlongitudinalmovement associated withspatialconnectivityinfreshwater this chapter, wehavediscussedhowthemainprocesses consideration whenplanningforfreshwater biodiversity. In ecosystems structure andfunction,soitneedsspecial Spatial connectivityisamajorcomponentoffreshwater conservation planninginthistraditionallyignored realm. excuses todelayoravoidimplementationofsystematic Biology, and variedapplicationsacross theglobe(e.g. conceptual understandingoffreshwater ecosystems— Given theadvances—intermsofmethods,dataand 9.4 Conclusions Threats andthesetofmanagementactionsto Thedataare more manageable. Thisaccuracyleadstoanincrease ineffi Attributeinformationismore accurate(asdiscussed management actions. of threats andthesubsequentspatialallocationof a reduction inuncertaintyforthespatialdistribution address themcanbetackledmore effi can hitthistarget more directly. cover atarget –say, 100habitatkmofaspecies,one in theparagraphabove). Vol. 56:1Jan.2011), there shouldbeno ciently are nolongerjustifi ad hoc ciently through orscoring Freshwater ciency. To ciency. ed by ner c CHAPTER 9 | FRESHWATER ECOSYSTEMS 287 icts and minimizing the cost of its icts and effectively. However, the implementation of such plans However, effectively. of the does not necessarily imply the strict protection whole catchment (e.g., see Section 9.2). Systematic conservation planning also accounts for the socio- economic component of conservation, by identifying way of maintaining the targeted the most cost-effective This enhances the likelihood of success for biodiversity. the implementation of conservation plans, by reducing potential social confl completion. © ROGER BILLS community from paying more attention to freshwater attention to freshwater paying more community from conservation planning, while marine and terrestrial developed. The traditional obstacles being plans were by the planning engendered conservation to freshwater misconception of planning and implementation scales of times by the proposal in recent has been overcome mixed conservation-land uses schemes. Freshwater conservation should ideally be planned at the whole catchment scale (minimum functional unit), since this more of ecological processes allows the addressing Lower Orange river at Pella Drift. Chapter 10. Development and policy plans for the future

Cross, K.1, Niskanen, L.1 and Dragisic, C.2 (Section 10.1); Barrera, L.3 and Canter Weikel, M.4 (Section 10.2)

1 Eastern and Southern Africa Regional Ofﬁce (ESARO), IUCN 2 Center for Environmental Leadership in Business, Conservation International 3 Center for Conservation and Government, Conservation International 4 Center for Environmental Leadership in Business, Conservation International Improper siting and management of agricultural development can lead to increased sedimentation, pollution and eutrophication of wetlands. Agriculture is also responsible for approximately 70% of water withdrawals from freshwater systems. © JEFF ATTAWAY

CONTENTS 10.1 Wetland development plans for Africa 290 10.1.1 Wetland resources and uses 290 10.1.2 Degradation and loss of Africa’s wetlands 291 10.1.3 Development across Africa 292 10.1.4 Impact of wetland development on inland waters 297 10.1.5 Threat mitigation and management plans that integrate freshwater biodiversity 297 10.2 Freshwater biodiversity management and conservation, policy, and the private sector 299 10.2.1 The policy process 299 10.2.2 Role of the private sector 302 10.2.3 Policy and the private sector – conclusion 305 290 CHAPTER 10 | DEVELOPMENT AND POLICY Nile (50,000km 20,471km Africa designatedundertheRamsarConventioncovering As ofDecember2010,there were 298sitesinmainland estuaries andmangrove swamps. floodplains andtheOkavangoDelta,aswellcoastal Africa,theresouthern are the ZambeziandLimpopo Cameroonnorthern (Wetlands 2007).In International NigeriaandtheChari Logonefloodplainsin in northern the SenegalRiverDelta,Hadejia-Nguruwetlands Moser 1991).Othermajorwetlandsacross Africainclude the Congoswamps(80,000km three ofthelargest wetlandsystemsinthecontinent, (Finlayson andMoser1991).EquatorialAfricacontains but are stillofgreat importancetopeopleandwildlife smaller orseasonalwetlandswhichhavenotbeenmapped, contains wetlands,andthisdoesnotincludethemany About 1%oftheAfricancontinentsurface(345,000km 10.1.1 Wetland resources anduses 10.1 © ART WOLFE/WWW.ARTWOLFE.COM Lake NarashainKenya,partoftheVictoria basinwhichisoneofthelargestwetlandsystemsinAfrica. cycles for fishing,livestock grazingandrecession agriculture. many livelihoods withinfloodplainsdependonseasonal flood as supplementary incomeformillionsofpeople.For example, immediate benefitsintermsofwater andfoodsecurity, aswell see Sections1.3and8.4,thisvolume). Wetlands alsoprovide groundwater throughout theyear (Bakema in spaceandtime,toensure availabilityofsurfaceand and servicesincludingthestorage anddistributionofwater African wetlandsare crucialto providing ecologicalgoods in Africa(Figure 10.1). Victoria Basin(approximately 50,000km Africa Wetland developmentplansfor 2 , whichisnearly6%ofthetotalarea ofwetlands 2 ), andmultitudeofwetlandsintheLake 2 ), theSuddinUpper 2 ) (Finlaysonand et al . 2009) (also . 2009)(also 2 ) So, and sandmining, andcraftmaterials(Bakema fuel woodcollection, dryseasongrazing,water supply, clay bottoms are usedforfoodproduction, fishingandhunting, Lake edgewetlandsandmany seasonal wetlandsinvalley Source: Wetlands 2011. International covering nearly6%ofwetlandsinAfrica. Convention haddesignated298sitesacross Africa, Figure 10.1ByDecember2010theRamsar unless wetland ecosystems are managed appropriately, et al . 2009). . 2009). CHAPTER 10 | DEVELOPMENT AND POLICY 291 . 2007, in et al © ROBIN MOORE ) and the water hyacinth Lates niloticus ), agricultural runoff, and overfishing of ), agricultural runoff, . 2010). In Lake Victoria, the introduction of the the introduction . 2010). In Lake Victoria, et al Eichornia crassipes Thieme ( invasive Nile perch and habitat native species, combined with climate change (and certainly degradation, have caused the possible loss of the cichlid proportion decline in abundance) of a large drinking water, irrigation for commercial and subsistence and for commercial irrigation water, drinking as serve They fishing grounds. and hydropower, crops, and sewage systems. venues, transit corridors recreational can ecosystems and other freshwater While wetlands overuse or improper impacts of these activities, absorb the by degradation caused can lead to ecosystem management nutrient sedimentation, turbidity, flows, increased changes in organic oxygen content, increased enrichment, decreased the entrance of invasive species,content, pollution, and so are there why Part of the reason among other effects. because national ecosystems is to freshwater many threats of the unaware and internationalmakers are level decision in their values of the wetland resource ecological needs and matters, complicate (Finalyson and Spier 1999). To charge often have cumulative impacts, making the diverse threats its individual parts and mitigationthe sum worse than complex. strategies more resources The impact of development on freshwater withdrawals and be seen in Africa. Water can already and Lake Adele sedimentation have caused Lake Haromaya in Ethiopia to dry out completely (Alemayehu ( © TIMO MORTIZ 10.1.2 Degradation and loss of Africa’s wetlands 10.1.2 Degradation and loss of Africa’s much of human settlement has occurred history, Throughout sea coasts, andalong rivers, oases, and lakes, as well as the use of waterhuman development is intimately tied to of the world, freshwater as in the rest In Africa, resources. with ecosystems supply cities and rural communities the functions that support agriculture, other food security the functions that support agriculture, undermined. and ecosystem services are In Africa, as in the rest of the world, freshwater ecosystems supply cities and rural communities with drinking water. In Africa, as in the rest of the world, freshwater supply cities and rural communities with drinking water. ecosystems A fisherman Faso. cast netting in a wetland in Burkina income for millions of provide food security and Wetlands people across Africa. 292 CHAPTER 10 | DEVELOPMENT AND POLICY capita, a proxy forincome, isjustUSD 1,127withwide rate of2.5%per year. Gross DomesticProduct (GDP)per stood at840million in2009,isexpectedto grow ata Africa isundisputed.Thecontinent’s population,which The needforeffective, sustainabledevelopmentacross appropriate mitigationplansare notputinplace. development planscouldwell increase thisfigure if (see Chapters3-8,thisvolume).Africa’s ambitious species assessedfortheIUCNRedListare threatened It isalready estimatedthat,in Africa, 21%offreshwater discussed above)islikelytobecorrespondingly large. other freshwater bodies,and freshwater biodiversity(as African continentissignificant,andtheimpactonwetlands, The scaleofdevelopmentunderway, andproposed, onthe 10.1.3 Developmentacross Africa industrial uses(IUCNESARO,unpublished). wetlands havebeenclaimedandconvertedtourban functions ofthesewetlands have resulted inthelossesofsomehydrological and Barotse floodplainshascausedhabitatchangesthat Rufiji (Tanzania), andShire River(southofLakeMalawi) The harvestingofgrassesandreeds intheCaprivi/Chobe, reeds andgrasses,isthreatening wetlandecosystems. mounds. Overexploitation ofriverinevegetation,suchas banks ofponds,oxbowlakesandlagoons,ontermite floodplain (Zambia),peopleare settledontheriverlevees, pressure onthewetlandresources. IntheBarotse people reside inthewetlandareas, exertingimmense In theCaprivi(Namibia),recent estimatesare that83,000 this hasresulted inhabitatchangeandlossofspecies. population pressure (from bothpeopleandanimals), Furthermore, there are water infrastructure overthelast80years(Dugan1990). population increase andpressures suchasdamsandother loss anddegradationofnaturalecosystemsdueto andcentralAfrica, thereIn western hasbeensubstantial At alocallevel,many catchment inTunisia (FinlaysonandSpier1999). of 15%wetlandarea and84%lossintheMedjerdah Other studiesreport thatthere hasbeenanoverallloss the Mfolozicatchmentwasestimatedtohavebeenlost. Tugela Basin(inNatal),and58%oftheoriginalwetlandin over 90%ofwetlandresources havebeenlostinthe wetland lossismuchgreater. Forexample,inSouthAfrica is around 2%.Incertainareas, however, itisknownthat that overallwetlandlossintropical andsubtropical Africa the 20thcentury. Values givenbyOECD(1996)suggest areas across theglobedecreased inextentby50%during other uses.According toWWF, itisestimatedthatwetland Wetlands are alsooftenlostcompletelywhenconvertedto al species nativetothelake(Witte . 2008;Hecky et al . 2010;allinThieme wetlandsinAfricaare experiencing estimatesthathalfoftheworld’s (IUCN ESARO,unpublished) et al . 2007;Chapman et al . 2010). et . South Africa and northern Africa,which areSouth Africa andnorthern water-stressed. materials. Thisis, bydefinition,aproblem inareas suchas as waterisoften usedforextractionandseparation of however, miningprocesses are extremely waterintensive, Mining practicesvarybyplace and commodity. Ingeneral, major multinationalcompanies. individual operationstolarge, commercial minesrunby Mines rangefrom small,informal (andoftendangerous) its platinumgroup metals(Smuts 2010;Mbendi2011). 40% oftheworld’s gold,60%ofitscobaltand90% 30% oftheplanet’s totalmineralreserves, including manganese, andbauxite.Africacontainsapproximately ranging from goldanddiamonds touranium,coal, continent. More than60products are minedinAfrica, Mining isamajor, andgrowing, industryontheAfrican Mining briefly below:mining,agriculture, andinfrastructure. freshwater systemsthroughout thecontinentare discussed three majorindustriesthathavesignificantimpactson biodiversity, willvarygreatly from placetoplace.However, ensuing impactsonfreshwater ecosystemsandfreshwater development contextforeachcountryorregion, andthe Africa isanextremely large andcomplexcontinent;the behind fewbenefitsforthelocalpopulation. production ofcommoditiesthatwillbeexported,leaving services), itrisksscarce resources beingusedforthe all, itmaybeseenasatypeofpaymentforenvironmental may bringinmuchneededincomeforAfricannations(after (TheEconomist2009). Whilethistypeofdeal governments set asideroughly onefifthofthecultivatedlandforArab official claimed thatSudanwould Sudanese government with majorGulfandAsianstates.Injustoneexample,a Africa Report2010).Thisincludeslarge-scale landdeals seeking tosecure accesstotheseresources inAfrica(The even scarcer, foreignare companies andevengovernments gain. Asavailablelandbecomesscarce, andwaterperhaps and are seekingtodeveloptheseresources fortheirown seeing opportunitiesintherichnessofnaturalresources, At thesametime,actorsinotherpartsofworldare ambitious developmentplans. institutions are strivingtoaddress theseneedsthrough andother distributed tolocalpopulations.Governments be sustainedoverthelongterm,andwhosebenefitsare life expectancy, there isaneedfordevelopmentwhichcan income inequality, decrease infantmortalityandincrease statistics are farworse.To increase incomesandreduce Indicators, 2009).Inmanycountriesintheregion, the at birthisjust52years(World Bank,World Development mortality is81per1,000livebirths,andlifeexpectancy is unabletomaintaincurrent standards ofliving.Infant year, whichislowerthantherateofpopulationgrowth, disparities withintheregion. GDPgrowth of1.7%per CHAPTER 10 | DEVELOPMENT AND POLICY 293

© KEVIN WALSH adoption of best practices throughout the mining industry, the mining industry, adoption of best practices throughout (Smuts 2010). recommendations as well as to reﬁne in Africa. South Africa, widespread Mining is relatively and D. R. Congo Zambia Ghana, Zimbabwe, Tanzania, like Angola, Sierra all major players. In other countries are Leone, Namibia, Zambia and Botswana, mining is a earnings. currency The footprint of foreign major source in the near term. Demand of the industry will likely grow for new especially those required for mineral resources, technologies like cell phones and computer chips, is or are opened recently, Major new mines have increasing. under development, in South Africa, Namibia, Botswana, discoveries of (Mbendi 2011). Major and Tanzania in Mauritania and potential potential diamond resources southern Namibia marine diamond deposits in offshore the impact of increase (Mbendi 2011) could also greatly lakes, wetlands, and the the mining industry on streams, aquatic species that depend on these habitats unless the expansion is very well designed and managed. environmental (and social) impacts of large-scale mines mines (and social) impacts of large-scale environmental lag behind on (small-scale mining operations often development, sustainability issues). Best practices for siting, the negative effects reduce and management can greatly ecosystems and species (Sweeting of mines on freshwater the application and Clark 2000; ICMM 2006.) Lessons from foster further of these practices have been used to

An asbestos mine in Zimbabwe. Mining is widespread but can across parts of Africa providing income for national economies, also pollute connected wetlands impacting freshwater biodiversity. the use of water for mining may compete In these regions and water with alternative uses, such as agriculture, water of reclamation shortages may ultimately require (Hanlon 2010). Contamination of mining by-products from is a major concern mining processes for the water from example, results Acid mine drainage (AMD), for industry. come into contact with water sulfides in ore when iron not only makes the water sulfuric acid. This and produce highly acid, but also leaves behind toxic metals, such manganese, aluminum, nickel, iron, as arsenic, mercury, and uranium (Hanlon 2010). Other contaminants, such or the washing process as cyanide, enter the water from other mine waste if these the leaching of tailings and from such Needless to say, not well managed. materials are ecosystems contamination is devastating for freshwater and the species that depend on them. Like many of the both during operations impacts of mining, AMD is a threat and after a mine has closed, as the mine fills with water of a the removal (Hanlon 2010). Mining may also require disturbing water substantial quantity of topsoil and rock, systems, drainages, and flows. However, Mining will never be a low-impact industry. advances by the sector itself, as well as improved regulations in the host countries and the scrutiny of in the host countries regulations have led to substantial the international community, advancements in minimizing and mitigating the 294 CHAPTER 10 | DEVELOPMENT AND POLICY relative availability ofland, sunlight,cheap labour, and areas At thesametime, manycompaniesare realizing thatthe crop diversification,andequitygenderissues. a greater focusonenvironmental andsocialsustainability, seeking toinstilnewdevelopment paradigmsthatinclude the Green RevolutioninAsiaandLatinAmerica,are Many ofthesedonorshavereflected onthelessonsof such astheGatesFoundation(Gates2009). policy frameworkshavebeenannouncedbymajordonors improvement, small-holdersupport, infrastructure, and (or reintroduction) ofnewcrop varieties,technological development inAfrica.Investmentresearch, development of developmentinstitutionstofocusonagricultural Because ofthis,there isarenewed effort onthepart Foundation 2009). andAfricaisanetimporteroffood(Gates undernourished, expectancy. Oneinthree peopleinSub-SaharanAfricais to thecontinent’s highinfantmortalityrateandlowlife Africa thaninanyotherregion (FAO 2010),contributing percentage peopleremains ofundernourished higherin eminently suitableforcommodityproduction. Yet the scattered throughout. InmuchofAfrica,theclimateis production inthearidnorth,andsubsistenceagriculture growing forestry sectorinMozambique,shiftinglivestock ranching, andtimberproduction inSouthAfrica,a in Liberia,sophisticatedcommercial-scale agriculture, as thecontinentitself–colonial-erarubberplantations Agriculture andforestry production inAfricaisasvaried Agriculture andforestry and changingwaterflowregimes. sedimentloadsinrivers biodiversitybyincreasing inLiberia,canhavemajorimpactsuponfreshwater seenhere Deforestation, © CI/PHOTOBYJOHNMARTIN and competition with other water uses. and competition withother wateruses. increase, leading tofurtherproblems withover-abstraction theneedforirrigation could to modifyrainfall patterns, could leadtoaquifercollapse.As climatechangethreatens ecosystems andotherhumanuses and,ifextreme enough, for crop production candecrease wateravailablefornatural if theuseofirrigationincreases. Over-abstraction ofwater Thieme of freshwater withdrawals(Shiklomanov 2000;FAO 2009;in irrigated agriculture isresponsible forapproximately 70% eutrophication, andotherwater qualityissues.Globally, use, ifunblockedbyriparianbuffers, canleadtopollution, Run-off from improperly-managed fertilizerandpesticide can effect evapo-transpiration cyclesandwaterflows. Clearing ofvegetationcover, especiallyatalarge scale, development canleadtoerosion andsedimentation. biodiversity. Improper siting andmanagementofagricultural major impactonwetlands,otherwaterbodies,andfreshwater Agriculture andforestry production inanyformcanhavea are beingcontemplatedatlarge scales. Wheat, forest plantations,andjatropha, amongothercrops, on twomillionhectares inZambia(TheEconomist2009.) 2009). Chinaisreportedly alsonegotiatingtogrow biofuels production inD.R.Congo(Mongabay2009;TheEconomist develop betweenoneandthree millionhectares ofoilpalm one example,aChinesefirmhasannouncedplansto development plansatascaleseldomseenbefore. In in co-ordinationare withnationalgovernments, announcing regions. Large multinationalcompanies,sometimesworking competitive advantageincommodityproduction overother abundant inwaterandgoodsoilspartsofAfricaprovide a et al . 2010),anumberlikelytobemirrored inAfrica

CHAPTER 10 | DEVELOPMENT AND POLICY 295 he According to the FAO (2007), to the FAO According he Africa Water Vision (AWV) 2025, developed in 2000, (AWV) Vision he Africa Water Dams and water related infrastructure related and water Dams has Africa currently infrastructure, large-scale In terms of of measures control dams and other water the fewest in the world. any continent Despite the low level of water infrastructure compared to compared Despite the low level of water infrastructure in recent pressure been increasing has there other regions, as a means of measures years to adopt such water control development. T enhancing food security and promoting which focuses on the crucial role of water in accomplishing which focuses on the crucial role goals. Targets the needed socio-economic development of safe and adequate water supply include: the provision doubling the and sanitation for 95% of the population; and developing 25% of to 24 million hectares; irrigated area 2000). This vision needs (UN Water/Africa the hydropower Resources was endorsed by African Ministers of Water Forum in Water and other stakeholders at the 2nd World Forum in Istanbul in Water May 2000. At the 5th World to financing requirements 2009, an update on the March The estimated investment achieve this vision was provided. under the AWV water infrastructure needed for Africa’s was USD 20 billion per annum at the time it was prepared in 2000. An updated estimate in 2008 puts the required investment at USD 50 billion per annum. The principal in estimations lies in the near ten-fold increase difference to achieve infrastructure of the cost of water resources securities and hazard food and energy economic growth, management, with the estimate for the development of multi-purpose storage power (HEP) driven hydroelectric the dominant) probably is part of the mix and (hydropower put at USD 20 billion per annum. and multiple uses are A number of dams for hydropower either under construction or being planned on the African there are 1,300 large- and medium-size dams in Africa, dams in Africa, and medium-size large- 1,300 are there also Chapter 1, South Africa (see in are 40% of which The majority of these have been constructed 1.3). Figure deal with rising demands for water in the past 30 years to of dams have populations. The majority growing from to water for irrigation (52%) and been built to provide of percent (20%). Twenty supply water for municipalities for multiple use, which nearly always includes dams are fact that only 6% of dams were irrigation. Despite the as a primary function, hydroelectric built with hydropower power than 80% of total for more power (HEP) accounts World to the countries. According generation in 18 African dams have been Commission on Dams, only 1% of African 2007). (FAO flooding control constructed to provide development of well planned infrastructure is recognised is recognised development of well planned infrastructure especially in as a critical component of economic growth, including such infrastructure, Sub-Saharan Africa, where This includes dams, is built for a variety of reasons. of both domestic and agricultural water supplies provision floods and control power, to provide water is scarce, where emphasised in (McCartney and Sally 2005). This need is t © JEFF ATTAWAY As with mining, many of the potential negative impacts As with mining, many of the potential negative ecosystems on freshwater of agricultural production siting and proper and species can be mitigated through voluntary management decisions. In the case of agriculture, adherence may require and donor requirements standards For to stricter guidelines than government regulations. Palm Oil (www. example, the Roundtable on Sustainable on Sustainable Biofuels (http:// and Roundtable rspo.org) as do include criteria on freshwater, rsb.epfl.ch/) standards Policies (www.worldbank.org). Safeguard Bank’s the World siting; drip Best management practices include: proper fertilizer irrigation (when irrigation is necessary); precision and maintenance of soil application; no-till agriculture; widely known and riparian zones. Many of these are cover, This does not mean educated producers. among more among less compliance is universal; it is likely to be lower sophisticated companies and less educated producers, weak or enforcement are regulations and lower still where the potential negative lax. Still, the means exist to reduce on freshwater. impact of agricultural production and forestry This is especially important as agriculture to the needs of a in Africa, in response continue to grow of outside players. population and the demands growing domestic in improving With investments being promised Africans, for the poorest especially agricultural production, and sophisticated commercial and the entrance of large be at a turning may investors, African agriculture point. If Africa’s agricultural development is managed intelligently, the underpinnings of may provide land and water resources If not, unsustainable agricultural growth. sustained future development — especially at the scale of expansion being discussed — could be highly detrimental to the continent’s and could undermine ecosystems and species, freshwater options for Africa’s development the sustainability of future people. Improper siting and management of agricultural developmentImproper siting and management can lead to increased sedimentation, pollution and Agricultureeutrophication of wetlands. is also responsible freshwaterfor approximately 70% of water withdrawals from systems. 296 CHAPTER 10 | DEVELOPMENT AND POLICY Section 1.2.2.1). InCameroon, there are developmentplans produce upto39,000MW ofelectricity(butsee Chapter1, proposed forthe CongoRiverinD.R.Congo, andcould Grand Inga,theworld’s largest hydropower scheme,is of theCongoRiver, hasrecently beencommissioned. Congo, theImboulouDamon the LefiniRiver, atributary to BurkinaFaso,Côted’Ivoire andMali.IntheRepublicof dam hasthepotentialtogenerate enoughpowerforexport being constructedatBuitoimprove energy security. Thenew under construction.InGhana,ahydropower damiscurrently 2,600MW MambillaDamhasbeenapproved butisnotyet Boa MariaDamonthePungueRiver. InNigeria,the downstream ofCahoraBassa. There isalsotheproposed Nkuwa project ontheZambezi Riverhasbeenapproved In Mozambique,constructionofthe1,500MWMphanda are alsoplansforGibeIV(1,472MW) andGibeV(560MW). III project and100MWFinchaAmerti NesheProject. There construction inEthiopiaincludethe1,870MWGilgel-Gibe been builttogenerate300MWofelectricity. Damsunder region. InEthiopia,theTekeze Dam,completedin2009,has 600 million.Zambiaplanstoexportmore electricitytothe capacity ofabout750MWandanestimatedcostUSD Lower KafueGorge Dam,whichplanstohaveagenerating built ontheNileinSudan.InZambia,there istheproposed Merowe DamisaUSD1.8billionhydropower project being continent. Anumberoftheseare describedbelow. The neglected. development,howeverifthisdevelopmentistobesustainabletheir impactsonecosystemsandbiodiversitycannotbe future The AkosomboDam,ontheVolta River, Ghana.The constructionofdamshasbeenidentifiedasamajorpartAfrica’s © KEVINSMITH to study properly theenvironmental and socialimpact and neglected. the impactson ecosystemsandbiodiversity cannotbe If waterandwetlanddevelopment istobesustainable, countries inAfrica(FAO 2007). larger than1,000hectares existinabouttwothirds of Madagascar, Morocco, South AfricaandSudan.Schemes 13.4 millionhectares, ofwhich nearly70%isinEgypt, total area underirrigationinAfrica isestimatedtobeabout The Development Bank,aswellnationalgovernments. that are oftensupportedbyagencies suchastheAfrican are alsonumerous irrigationprojects throughout Africa or watersupplyfordomesticuseand/orirrigation.There There are alsomanysmallerdamprojects formultipleuse and DamConstruction2011). Water Rivers2011;International 2011 (International Power Hydroelectric PowerPlantthatisexpectedtobeginearly been completed,andconstructionofthe700MWKaruma Nile River, whichwasapproved in2007andhasnotyet 250MW power-generating facilitybeingbuiltontheVictoria include theBujagaliHydropower Project (HPP)whichisa and Namibiaare underway. InUganda,twonewdams for theBaynesDamonKuneneRiverbetweenAngola for LomPangar, Memve’eleandNachtigaldams.Studies There isanunderstanding thatitisimperative CHAPTER 10 | DEVELOPMENT AND POLICY 297 Maintenance © ABEBE GETAHUN Threat mitigation and management plans that Threat biodiversity integrate freshwater major ecological and environmental changes in in changes environmental and ecological major resources natural decreasing in resulting floodplains, biodiversity and fish stocks, pasturelands such as International Delta, In the Inner Niger 2007). (Wetlands governing of use and management the the process over has deteriorated including water natural resources governanceto natural, socio-economic, time due and the government Consequently, management constraints. Niger degraded habitats in the Inner of Mali is restoring of local people, decentralized Delta with the involvement government, partners in the Delta areas. and development to the national environmental This is contributing directly International 2007). policy (Wetlands protection supplies, declining water quality Diminishing water wetland habitats due to upstream and conversion of all have knock- encroachment impoundment or local few long-term are While there on biodiversity. on effects a number of studies biodiversity monitoring programmes, biodiversity in have noted dramatic declines in freshwater within historical time scales and Victoria Lakes Tanganyika (IUCN ESARO, unpublished). 10.1.5 Africa, across Although wetlands face serious threats water governance institutions that can manage wetlands a potential mechanism for for multiple use present sustainable management of this ecosystem. following 2 from an original area of 3,000km original area an from 2 Impact of wetland development on inland wetland development Impact of waters Lake Wenchi, a crater lake in Ethiopia. The Wenchi Eco-Tourism Association (WETA) is a local, community-centred is initiative Association (WETA) Eco-Tourism a crater lake in Ethiopia. The Wenchi Lake Wenchi, which promotes sustainable eco-tourism to improve the livelihoods of local people. Water infrastructure can have a variety of negative impacts can have a variety of negative impacts infrastructure Water peaks flood due to reduced on wetlands. For example, of overbank and extent be a lower frequency can there in the Hadejia-Nguru wetlands flooding. For example, than to less was reduced in Nigeria, annual flooding 1,000km provide proper compensation to all those communities those communities to all compensation proper provide many However, affected. are that environment and the wetlands. will be impacted, including ecosystems 10.1.4 construction of upstream dams. Dams can result in a dams. Dams can result construction of upstream load in rivers, in sedimentation reduction downstream and beds, of river-banks erosion leading to increased and degradation of coastal deltas. loss of floodplains on sediments wetland systems that rely This can affect often and flooding to sustain vegetation. Fish populations areas. supported by wetlands, especially as breeding are impacts on Destruction of wetlands can have significant such as dams infrastructure fish populations; furthermore, and disconnect rivers from can block migratory routes, Reduced their floodplains (and associated wetlands). vegetation that flooding that supports wetlands can alter and bird may be important for a wide range of mammal species (McCartney and Sally 2005). systems and various other types of Flood control have caused development in the Sahelian river basins 298 CHAPTER 10 | DEVELOPMENT AND POLICY strategies formulatedwhichincorporatethestakeholders. wetlands outsideprotected areas, andnewmanagement component species,greater efforts shouldbefocusedon to beconservationofthreatened andfragilesitestheir Although thegoalforprotected wetlandsshouldcontinue by wetlands. that takesintoaccounttheecosystemservicesprovided policies andcreating asustainabledevelopmentframework to adoptinganIWRMapproach inallwatermanagement and CentralAfrica) (Westand managementofwetlandsinfourecoregions from thereport managing theworld’s waterresources. Recommendations increasingly recognised inthewide-rangingdebateon management ofwetlandsandtheirbiodiversityisbeing their management.Indeed,theimportanceofsustainable policies andencouragementforcommunityparticipationin political willtoprotect them,basedonsoundwetland The future oftheregion’s wetlandsdependsuponastrong (Finlayson andMoser1991). irrigation schemesthatcanincorporatewildlifesanctuaries smaller damsacross rivers,andsmallermore diverse from damstoprovide environmental flows,constructionof infrastructure’. Interventionscanincludecontrolled release and promoting theconceptthatwetlandsare ‘natural water andwetlands,awareness raisingandeducation, This requires strengthening institutionsthatmanage as theneedsofindustrialandcommercial development. including people,wildlife,ecosystemdynamics,aswell (IWRM), whichtakeintoaccountallaspectsofwetlanduse principles ofintegratedwaterresource management biodiversity theysupportisthrough implementingthe areas and offers anexample ofstrong politicalwill to encompasses wetlands inprotected andnon-protected for themanagement ofwetlands.Thewetland policy departmentresponsiblein Africathathasagovernment Resources in1995,anditisoneofthefewgovernments Policy fortheConservationand ManagementofWetland 2009). policies andstrategiesatthenational level(Bakema wetlands isthrough thestrengthening andintegrationof A keystepinachievingsustainablemultiple-useof The conservation measures (Bakema these broader benefitsare more likelytoraisesupportfor a hydrological functionorcanpromote tourism,then to keyresources. However, ifconservationefforts have of supportbylocalpopulationswhomayloseaccess with noprovisionuse,itcanresult foralternative inalack management focusespurely onbiodiversityconservation their livelihoodsstrategies(Bakema people tousewetlandresources asakeycomponentin an unrealistic optioninAfrica,asthere isareal needfor of wetlandstosolelyprotect biodiversityis,inmostcases, way forward tosustainingwetlandsandthe The government ofUgandapassedtheNational The government Strengthening policiesforthewiseuse Strengthening byWetlands (2007)point International et al. et al 2009). . 2009).Ifwetland et al . and institutionalcapacity(IUCNESARO,unpublished). limited infrastructure development,sustainablefinancing for effective implementationofIWRM.However, there is the necessarypolicy, legalandinstitutionalframeworks the enablingenvironment. Mostcountrieshavedeveloped institutional capacityprovide ayardstick formeasuring of sustainablefinance,institutionalarrangements,and legislation, levelofinfrastructure development,availability enabling environment withinthecountry. Thepolicyand of IWRMprinciplesdependsontheexistence Wetland Inventory The and theirspeciesacross thecontinentmustbeincreased. For sustainablemanagement,theknowledgeofwetlands impact assessment. flows, inwaterresource allocationandenvironmental to takeintoaccountallwaterneeds,especiallyenvironmental To avoidfurtherenvironmental degradation,countriesneed unpublished). conserve wetlandsandtheirbiodiversity(IUCNESARO, multiple use. with otherknowledge systemsthatencourage sustainable the basisforwetland managementsystemsinconjunction benefits are alsoneeded.Localknowledgeshouldform different usersandguarantee equitabledistributionof Respected institutionsthatcan resolve conflictsbetween that spelloutrightsandresponsibilities ofallstakeholders. According toBakema in placeandfunctioning. decisions, institutionalframeworksandcapacityneedtobe to usetheinformationmakeinformedwaterallocation water resources are essentiallyalready fullyused.Inorder wetlands, andtoadecrease inHEP generationasthebasin’s decline inthehealthofriverecosystemincluding maximised agriculturaldevelopmentwouldleadtoafurther some lossofagriculturalarea. Conversely, scenariosthat wetland withnooveralldecrease inHEPgenerationbutwith river conditionandsubstantialre-flooding ofadownstream rules fortheHEPdamscouldallowsomeimprovement in scenarios foundthatabasin-widerevision oftheoperating an in-depthflowassessmentwhichdevelopedanumberof Sally 2005).Forexample,inthePanganibasinTanzania, impacts ofdifferent flowrelease strategies(McCartneyand the complexprocesses andinteractionofflowsthe scientific andengineeringstudiesare required tounderstand and otherdownstream ecosystems.However, detailed from damscancontributetotherestoration ofwetlands the statusofwetlandsinAfrica.Environmental flowreleases further research inorder tohaveaclearer understandingon and climate(FAO 1998).Therefore, there isacriticalneedfor such ashydrology, socio-economy, soils,land-use,vegetation be collectedonbiophysicalandsocio-economicparameters quantitative studiesonwetlandlossinAfrica.Informationmust like SouthAmerica,there isanextreme lackofpublished Global Review of Wetland Resources and Priorities for Global ReviewofWetland ResourcesandPrioritiesfor (Finlayson and Spier 1999) found that, (FinlaysonandSpier1999)foundthat, Adoption andeffective implementation et al . (2009),wetlandsrequire rules CHAPTER 10 | DEVELOPMENT AND POLICY 299 nowledge exchange . 2003). Once the agenda is set, then change et al Freshwater biodiversity Freshwater management and conservation, and the private sector policy, 10.2 of freshwater Successful management and conservation of a variety biodiversity necessitates the involvement including local communities, of stakeholder groups, The role sector. scientists, policy makers, and the private makers and the private of the latter two groups—policy detail below. in more sector—is explored 10.2.1 The policy process an ongoing Policy making at all levels is essentially change that is punctuated by of incremental process opens up. times of rapid change when a ‘policy window’ is an essential Good information, based on good data, both from component of ensuring that policies emerging sound. are of these phases of the process policy solution, a that requires is always a problem There of decision makers and the values, beliefs and perceptions shaping the problem, this influence attitudes towards for negotiation. Coalitions as solutions ideas presented governmentof stakeholders from institutions, advocacy the general public, the private sector and organizations, beliefs and interests core that share the political process perspectives to the policy their come together to promote of those coalitions will The most dominant problem. have the upper hand in terms of setting the agenda for that serve their interests negotiation and ensuring results (Johnson change’, tends to be slow; this is the period of ‘incremental and change during this time is accomplished by influencing those basic values and beliefs of the stakeholders. A key the level of knowledge and way to do this is by increasing understanding of the stakeholders so that, as their views policy and beliefs change over time, so do their preferred solutions; that is, it is a period of gradual ‘enlightenment‘ based on the accumulation of information (Sabatier 1991). downstream wetlands and cultivation areas. This resulted resulted This areas. cultivation and wetlands downstream wetlands. use of the multiple from loss of livelihoods in the in managing and restoring investment the current However, in clearing – wetlands – has resulted natural infrastructure of wetlands restoration for water transport, water channels support grazing of dams to and re-operation for fisheries been 2010). This has all on floodplains (Smith and farming actions: k made possible by several and capacity development for decision makers and and capacity development of governance arrangements for stakeholders; the reform accountability of government institutions, water; increased building among stakeholders; and enabling consensus a trust fund for basin restoration and establishment of land and water management, including and improved (Smith 2010). community-led action et . 2010 for discussion of the cost of engineered solutions . 2010 for discussion of the cost of engineered The water resources management strategies in the The water resources still very much guided by the traditional are region to water management, whose main emphasis approach of is on engineering solutions without consideration goods and services (see also Vörösmarty environmental al of freshwater and the implications to management aquifers ecosystems). The soils, wetlands, watercourses, as natural and floodplains of watersheds can be considered moves, stores, Just as built infrastructure infrastructure. Lakes and so does nature. and cleans water, regulates reduce water storage and can therefore wetlands provide volume needed in dams, consequently reducing reservoir and the associated cost. the need for built infrastructure infrastructure Thinking about wetlands in terms of natural a catchment with the concept of managing introduces the both built and natural components and reduces and conflict between engineering solutions perceived (Smith 2010). nature in sustaining natural infrastructure The investment required water infrastructure to that in engineered is different financing and top down which need significant projects, institutions and governments. large decision making from needed to Instead, bottom up strategies and actions are for multiple uses. What or conserve wetlands rehabilitate stakeholders with the is needed is to empower relevant and build consensus on capacity to negotiate trade-offs as people (Smith 2010). as well priorities that benefit nature For example, if part of a wetland is to be used for rice or regulation be agreement needs to cultivation, then there destruction and on how this will be managed to prevent loss of benefits to other users (including the environment). basin of northern Nigeria, dams In the Komadugu Yobe in damage to built in the 1960s and 1970s have resulted The balance of the local community’s activities around around activities community’s the local of The balance ecosystem of the part an essential use is wetland to protect areas conservation Therefore, dynamics. to maintain the balance must be designed wetlands the is through An example and nature. between people Community through Areas Protected Wetland Extending as COBWEB) in Uganda, Initiatives (known Conservation United Facility, the Global Environment funded through government Programme, of National Development The rationale is that wetland Uganda and other partners. conserved and the socio-economic biodiversity needs to be must be maintained of these areas importance and values in managing the protected by engaging the communities The goal is to establish and strengthen wetland areas. and sustainable wetlands community-based regulation biodiversity. use within wetlands with important resource by IUCN being implemented is currently The project it will have and partners, and by its conclusion in 2012 biodiversity community conservation models for wetland and processes integrated into national wetland planning network. area national protected 300 CHAPTER 10 | DEVELOPMENT AND POLICY study is an importantexample ofhowto integratescientific The Economics ofEcosystemsandBiodiversity (TEEB) building trustwithpartnersand otherstakeholders. Engagement throughout theprocess isalsoimportantfor slow periodsandthe ofrapidpolicychange. order tohavescientificinformationreflected bothinthe by engagingpolicyleadersand like-mindedgroups, in based. So,itiscrucialtobeinvolvedintheseslowperiods, (mentioned above)onwhichsubsequentdecisionsmaybe an importantpartoftheoverall’periodenlightenment‘ proposals hadnotbeenthedominant views,theyare still not beenthedominantviews.Evenifinformationand proposals thatare already onthetable,eveniftheyhave are, therefore, almostalwaysbasedoninformationand established ideologicalandcontextualframework is created, theensuingrapid changes occurwithinan policy windowsopenup.Thisisbecause,whenawindow are actuallyveryimportanttoachievingkeygoalswhen an unprofitable commitmentofresources. However, they slow, anditcanseemthatengaginginthemrepresents change thatcomebefore andafterthepolicywindowsare involved. Bycomparison,theperiodsofincremental everyone witharelated interest tendstowantbe in policyattractalarge numberofactorstotheprocess; and decisionsbeingmade,thehighratesofchange In timeswhenpolicywindowsopen,thenewinformation in thenextsection. context tothesupportingdata.We’ll examinethisfurther be basedonreliable data,buttheywillhaveadifferent new outcomes.Thisinformationandthesolutionsmust the opportunitiestomovethrough thiswindowandachieve information andpotentialsolutions,inorder tomaximize has beenopened,however, itisnecessarytoprovide good the policydiscussionintopubliceye.Oncewindow related totheissue;orothersimilareventsthatcanpropel emergence ofnewinformation;uncoveringcorruption development project withfarreaching implications;the of somesort,suchas:anaturaldisaster;plansforlarge to bringrenewed attentiontoanissue–thiscanbeacrisis policy windowisusuallycatalyzedwhensomethingoccurs proposal (Kingdon1984).Inpractice,theopeningofa circumstances thatmakeitadvantageoustosupportthe problem, aproposal foraviablesolution,andthepolitical of apolicywindowrequires three elements:aknown in policydesignare usuallyachieved.Thecreation occur inthepolicymakingprocess, andgreat leaps When a‘policywindow’opensup,rapidchangescan person canbethekeytocreating change. is criticaltoachievingchange;oftenjustonededicated and influentialchampionsthatare apartoftheprocess policy makingisapoliticalprocess, andhavingvocal cause ofinterest withinthelarger policyarena. Ultimately, influential individualswhocanthenchampionaparticular One importantstrategyistotarget oneorasmallnumberof to thesmall amountof floodingleftand wouldhave cut The construction ofthisdamwouldhaveput anend when afinaldam wasproposed intheearly 1990s. altered theamountofyearlyfloodingandwere inplace agriculture. Fivehydrodams on theriverhadalready which helpedtomaintainfertile riverbanksforseasonal streams, riverineforest andmangrove ecosystemsand regime whichsupportsgrasslands,lakes,seasonal Kenya’s hydropower. Theriverhasabiannualflooding Kenya. TheTana Riverprovides approximately 75%of ecosystem functionandvalueintheTana Riverbasinin Emerton (2005a,b)reported on someanalysesof Tana riverbasin Economic evaluationofhydropowerdevelopmentinthe here. community. Two examplesspecifictoAfrica,are given to theinterests ofstakeholdersbeyondtheconservation discussion, andtheydosoinawaythatismeaningful present informationontheimplicationsofpolicy Useful analyseswillshare twokeycharacteristics:they Brink inpress). and recommendations forpolicymakers(TEEB2010;ten economic data(e.g.,Kumar2010)andpresent guidelines example, inseveralreports thatsynthesizeecologicaland policy decisions.TEEBhasattemptedtoachievethis,for synthesized toprovide results thatcandirectly influence helpful topolicymakers.Thedatamustbeanalysedand into policycanattest,rawdataalonemaynotbeespecially As mostscientistswhohaveattemptedtoinsertscience Increasing knowledgeandunderstandinginvolves ii. Collectivedecisionmaking,asinapolicycontext, i. consideration: data forpolicyapplications,twokeyideasmeritfurther In consideringhowtopackagefreshwater biodiversity actions movingforward. fields ofscience,economicsandpolicytoenablepractical degradation, anddrawingtogetherexpertisefrom the the growing costsofbiodiversitylossandecosystem the globaleconomicbenefitsofbiodiversity, highlighting initiative,whichaimstodrawattention international ’Potsdam Initiative‘forbiodiversity).TEEBisamajor five majornewlyindustrialisingcountries(theso-called of theenvironment ministersoftheG8countriesand of biologicaldiversity’,madeattheMarch 2007meeting a studyon‘Theeconomicsignificanceofthegloballoss policy (TEEB2011).TEEBisfoundedonaproposal for data intothedecision-makingprocesses ofbusinessand more thansimplyproviding rawdata. of knowledgedecisionmakers; knowledge andreliability ofexperts,andinthelevels is strongly influencedbybeingabletotrustinthe CHAPTER 10 | DEVELOPMENT AND POLICY 301 system. However, these data would not have been enough system. However, design option of the Mutonga-Grand to change the preferred on humanFalls Dam – the additional analysis of impacts impacts was livelihoods and the economic costs of those policy choice. needed to encourage a different Moulouya River Basin river regulation Basin Agency IUCN, working with the Moulouya River developed a and the University of Oujda in Morocco, management of aquatic for the integrated pilot project biodiversity at the level of the Moulouya River catchment was to incorporate (IUCN 2011a). The aim of the project into the catchment requirements environmental little was known management plan. Prior to this project, the Moulouya Basin, species within about the freshwater for their values to people, was little appreciation and there Catchment management or their ecological requirements. at this time was primarily focused on the allocation of the close involvement needs. Through water to people’s of a wide range of stakeholders, and an intensive the on findings from campaign based awareness-raising basin-wide biodiversity assessment conducted through A significantly altered. this focus has been the project, scientific committee has now been formed to focus on the management, and aspects of catchment environmental an ‘Aquatic Biodiversity Monitoring Network’ has been species established to monitor the status of freshwater the catchment. throughout © JEAN-PIERRE BOUDOT most of the downstream ecosystem off from water flow, flow, water from ecosystem off most of the downstream than 1 million people impacting the livelihoods of more for humans and cattle, fertile water sources by reducing for riverbank farming (because the banks of the ground and fish in the region), the only arable land are Tana and the main areas catches in the wetlands, mangrove river channel. analyses for the Mutonga-Grand Falls Pre-construction design option for the dam dam showed that the favoured mechanisms because of would not include flood release the cost involved. An analysis was carried out, looking at of the dam, downstream flood-dependent area the entire and detailing what the impacts on these ecosystems would that the construction of a new dam. This showed be from River had already the Tana of dams on the upper reaches million in lost production USD 27 imposed approximately costs, and that the cost of building an additional dam cost of almost USD 20 would have a median present million. With this information, which could be easily fit into appraisal framework, the traditional economic the more to most economically viable design option was revealed costs and integrated economic be the one that considered could simulate the Tana’s mechanism that a flood release biannual flood regime. This analysis would not have been possible without detailed flow and on the Tana’s data on the ecosystems that rely models of the river as well as hydrological flooding regime, The Oued Za, a tributary of the Moulouya River. The ecological needs of freshwater is now incorporated into the species The Oued Za, a tributary of the Moulouya River. Moulouya catchment management plan. 302 CHAPTER 10 | DEVELOPMENT AND POLICY some of whichhavemore influencethan others. multiple stakeholder groups, allwithdifferent interest – that requires useful analysis,anditisanegotiation amongst incremental changepunctuatedbytimesofrapid In allofthesesituations,policy makingisaprocess of large institutions,suchasmultilateraldevelopment banks. of nationalandregional policymakingandevenwithin influence theprocesses describedhere applyatallscales The policyprocess andthetype ofinformationneededto Raisingawareness ofbiodiversityvaluesand 3 Speakingtotheinterests ofotherstakeholders. 2 Usingdatatocreate scenariosofimpactsfrom 1 the policyprocess: successful integrationoffreshwater biodiversitydatainto These twoexamplesillustratesomeimportantfactorsin on thebasins’freshwater biodiversity. awareness raising,andtheprovision ofnewinformation influenced through involvementofkeystakeholders,public Morocco. Thisprovides anexampleforhowpolicycanbe as anexampletotheotherbasinagenciesthroughout into catchmentplanninginthisway, anditisnowpresented is thefirstriverbasininMorocco tointegratebiodiversity the ecologicalneedsoffreshwater species.TheMoulouya plan nowincorporatesaspecificcomponenttoaddress The result isthattheMoulouyacatchmentmanagement influence policy. data onspeciesdiversityandtheirecologicalneedsto the powerofstakeholderinvolvementandusereal needs. the more comprehensive analysisdid. likely tochangethecourseofdecisioninway of biodiversityconservation,butwouldnothavebeen support from thosestakeholdersthatshare thevalues only onthelossofspecieswouldhavehadplenty achieving thedesired outcome.Anargument based net negativeeconomicimpactwasthekeyfactorin River’s biannualfloodingcyclewouldactuallyhavea viability ofthedam.ShowingthateliminatingTana development from electricitygenerationandfinancial this case,theotherstakeholdersvaluedeconomic provides aparticularlygoodexampleofthis.In of landresource management.TheTana Rivercase withtherequirementsintegrating biodiversityconcerns Both examplesdemonstratetheimportanceof the policydebate. when there are legitimateinterests from bothsidesof is animportantpartofthepolicyprocesses, especially different designsofthedam.Providing viablesolutions and howthatwouldimpactlocalpopulationsunder development wouldimpactdownstream ecosystems, scenarios were presented forhowthehydropower different policychoices. The Moulouyacatchmentcasedemonstrates IntheTana Rivercasestudy,

determined withinabroader regional or globalcontext. any speciesidentified withintheassessment area tobe data onspecies isthatitallowstheconservation valueof status offreshwater species.Akeyfeature oftheRedList if usedwisely, helptoreduce impactsandimprove the their vulnerabilitieswillbegreatly improved andwill, data sets.Inthisway, theknowledge ofthespeciesand processes canalsohelptobuildfreshwater biodiversity as usefulinformationforinputtotheEIAprocess itself,EIA 2005). Inadditiontothedatainexistingsetsserving EIA process (Energy andBiodiversityInitiative2003;IAIA on howbiodiversitycanmore explicitly bebuiltintothe and implementingEIAs.There isevengrowing literature publication canbeextremely helpful forthosescoping the dataandtoolspresented in previous chaptersofthis and ecosystemservices,suchasfreshwater services,and EIAs are increasingly takinganexplicitlookatbiodiversity impacts. measures, andevaluationofsignificanceresidual mitigationand impactmanagement of alternatives, issues andimpacts,baselineestablishment,assessment regulatory framework,andtypicallyincludeidentificationof made (IAIA1996).EIAsare oftencompelledbyacountry’s prior tomajordecisionsbeingtakenandcommitments social, andotherrelevant effects ofdevelopmentproposals predicting, evaluatingandmitigatingthebiophysical, Assessment (IAIA),EIAsare theprocess ofidentifying, AssociationofImpact As definedbytheInternational within aproposed locationfordevelopment. the privatesectorshouldevaluatebiologicalsensitivity Impact Assessments(ESIAs)are additionalpointsatwhich Impact Assessments(EIAs)orEnvironmental andSocial in practicethisoftendoesnothappen.Environmental other integratedland-useplanningprocesses. However, tools likeStrategicEnvironmental Assessments(SEAs)or sector developmentbegins.Thiscanbeachievedusing time thatpolicydecisionsare made,thatis,before private sensitivity ofacatchmentshouldbetakenintoaccountatthe so somethingneedstobedone.Ideally, thebiological many speciesare threatened bydevelopmentactions mainland Africa.Thisreveals significantproblems, inthat analysis ofthestatusfreshwater biodiversityacross As reported here, IUCNhascompletedanin-depth business decisionsthatalsobenefittheenvironment. necessary biodiversityinformationtomakesustainable therefore, aneedtosupplytheprivatesectorwithall natural resources, suchasfreshwater systems.There is, huge role toplayinthemanagementandconservationof as recognized byTEEB,businessandenterprisehavea into thatarea toimplementthoseactivities.Inthatrespect, to occurinanarea typicallyprecede privatesectorentry Policy decisionsthatallowparticulardevelopmentactivities 10.2.2 Roleoftheprivatesector CHAPTER 10 | DEVELOPMENT AND POLICY 303 © RODERIC B. MAST monitor long term water quality. monitor long term water quality. The Okavango Delta is home to countless freshwater the Delta management plant the use of bio-indicators to and terrestrial species. It is now an aim to include within 304 CHAPTER 10 | DEVELOPMENT AND POLICY Conservation organizations are becoming increasingly biodiversity related considerationsinbusiness practices. opportunities created through inclusionofecosystem and aimed atproviding practicalguidanceontheissuesand private sector, andhasdevelopedareport (TEEBinpress) has recognized theimportanceofproperly informingthe is accessibletopolicymakers(as discussedabove).TEEB important asensuringthedata are presented inawaythat their developmentdecisions.Thischallengeisjustas to companiesinarelevant and usableformatforguiding It isimportantthatbiodiversitydatasetsare presented protected areas, andworldheritagesites. on thepresence ofreserves, regional ornationalparksand sets thatwillalsobeimportant–forexample,information quantity, qualityandtiming).There willbeotherbroader data being investigated(characteristicsofflowintermswater or artisanal);andtheecohydrology ofthefreshwater system species are threatened); utilizationofthespecies(commercial are endemic;populationstatusofspeciesandhowmany diversity ofspeciespresent (howmanyspecies;how the typesofdatathatare required are: informationonthe project development.Inthecaseoffreshwater ecosystems, likely itisthatthesedatawillpositivelyinfluencedecisionson to companiesconsideringdevelopmentprojects, themore comprehensive biodiversitydatasetsare madeavailable Considering theseobservations,itisclearthatthesooner resources are determinedtobebiologicallysensitive. the project moveforward, evenifthearea’s freshwater and otherstakeholdersare already quitevestedinseeing been sunkintotheproject, socompanies,governments, means thatalarge portionoftheinvestmenthasalready late inaproject’s developmentlifecycle.Typically, this take placeinarigorous manner, theyoftenoccurrelatively shortcomings (Jay However, EIAsastheyare oftenimplementeddohave ofthelakesurface. tensofthousandshectares covered Water hyacinthhadmajorecologicalandeconomicimpactsuponLakeVictoria during the1990swheninvasiveplant et al . 2006),andevenwhentheydo © STEPHANMAGDALINSKI (http://www.ibatforbusiness.org), was launchedatthe IUCN The firstoutputof theIBAT vision,the risk assessmentprocesses. information from allfourorganizations toinformbiodiversity first time,decision-makerscan accesscriticalsitelevel Conservation MonitoringCentre (UNEP-WCMC).Forthe and UnitedNationsEnvironment Programme World (CI),IUCN, ConservationInternational International, by anallianceofglobalorganizations, includingBirdLife This visionforIBAT wasdeveloped andisbeingrealized national orregional development strategies. networks ofconservationpriorityareas whendesigning project cycle.IBAT canalsobeusedtotakeintoaccount subsequent datacollection,assessmentandplanninginthe still economicallyviable.IBAT canhelpinformandprioritize approaches orlocationsatatime whensuchchangesare projects,planning makesiteasiertoconsideralternative Access tothisinformationattheearlieststagesofproject the project reported oninthisvolume. the continentbasedonspeciesdatacollectedthrough for Africa,andwilleventuallyincludefreshwater KBAsfor practice standards. Thisincludesterrestrially definedKBAs of environmental safeguard policiesandindustrybest Areas (KBAs)—IBAT informsthepracticalimplementation conservation—such asprotected areas andKeyBiodiversity on bothprotected andunprotected highprioritysitesfor use ofcriticalbiodiversitydata.Byproviding information vision forimproved collection,update,managementand The IntegratedBiodiversityAssessmentTool represents a Integrated BiodiversityAssessmentTool (IBAT). tools andapproaches toachievethis.Onesuchtoolisthe accessible tobusinesses,andhavedevelopedvarious attuned totheneedmakebiodiversitydatamore easily IBAT forBusiness tool tool CHAPTER 10 | DEVELOPMENT AND POLICY 305 et 2010) et al. ineffective management ineffective surface area, and the catchment’s annual and the catchment’s surface area, 2 2006). et al. . 2008). Many sources indicate one major contributing indicate . 2008). Many sources al weed on the lake was the of the factor to the proliferation agricultural development. In upstream nutrient inputs from sector by the agriculture understanding this case, a greater downstream affecting were inputs as to how their upstream the water hyacinth and of biodiversity (i.e., proliferation declining fish stocks) may have led to better practices and of crisis. impact during this time a reduced 10.2.3 Policy and the private sector – conclusion important are that there these examples, it is clear Across and the the private sector, linkages between policy, biodiversity. management and conservation of freshwater scientific and conservation easily accessible the The more communities can make information to policy makers, as is the chance well as companies and industries, the greater that positive conservation outcomes can be achieved. gross annual economic product is in the order of USD of is in the order annual economic product gross fish catch was 3 to 4 billion. During the infestation, the in addition to hampering by a factor of 2 to 45%, reduced facilities (Vincent lake transportation and hydropower ecosystems found in the delta, also represents the second also represents in the delta, found ecosystems earner income diamond mining), (after for Botswana largest Domestic Product Gross 5% of the country’s contributing in northern 40% of employment (GDP) and Botswana (Turpie management inspecific aspect of to one In relation IUCN (in collaboration with Thethe Okavango Delta, Harry Institute formerly known as The Okavango Research conducted a Centre) Research Oppenheimer Okavango long-term monitoring of improving case study with the aim use of bio-indicators (IUCN 2011b). of water quality through of this study was that a ‘coffee- A key recommendation be made which would appeal to alltable’ style publication stakeholders and key decision-makers across the relevant book (Mendelsohn The resulting the region. the the wonderful diversity of species within vividly presents and economies, livelihoods delta, and its value to people’s appealing to ain a format that is easily accessible and of stakeholders and decision-makers. cross-section broad of charge, free The book has been made widely available, an effective and it is hoped will prove the region, throughout way to influence policy. There are many examples where where many examples are There of freshwater resources, and the species present, can and the species present, resources, of freshwater This sector. inhibit the economic success of the private promote may be especially noticeable when conditions examplethe expansion of invasive species. A well-known invasive species of the ecological and economic impact of in Lake Victoria is given by the invasive water hyacinth in eastern which, during the late 1990s, blanketed Africa, surface. Lake of the lake’s tens of thousands of hectares lake in the world, freshwater is the second largest Victoria at 68,800km , IBAT IBAT Egretta ) which Grus carunculatus Afrithelphusa monodosus positively contribute to filling in positively contribute ) in north-western Botswana, for example, 2 provides key decision-makers with access to with key decision-makers provides ) also classified as Vulnerable) and large mammals and large ) also classified as Vulnerable) is a vast inland freshwater wetland and home to countless wetland is a vast inland freshwater species, including charismatic and terrestrial freshwater species of tourism value, such as two globally threatened ( species (the wattled crane bird ( and the slaty egret is classified as Vulnerable, vinaceiqula including the African elephant (Mfundisi 2008). The tourism species and dependent on the freshwater largely industry, World Conservation Congress (WCC) in October 2008. in October (WCC) Congress Conservation World for Business in terms of their interest, on sites of special critical information can so that this information and its conservation, biodiversity and address processes inform decision-making be used to 2008). (The Economist biodiversity impacts any potential needs, coupled assessment of business was built on an IBAT to integrate these databases into awith innovative work In addition to supporting integration web-based interface. and planning science into project of the best conservation opportunities also provides IBAT management decisions, to financially support the acquisition for the private sector and data on biodiversity andof additional knowledge including freshwater. conservation priorities, Companies can also freshwater data gaps through support for biological support data gaps through freshwater tools like IBAT surveys. When desktop surveys and use of that significant data gaps exist within the context of reveal under consideration, a biological a development project conducted prior to a formal survey of key taxonomic groups helpful in guiding early decision- ESIA can be extremely making for a potential development. developed by Conservation International,One such approach Assessmentutilizing the in-house expertise of the Rapid (RAP; also see Chapter 8, section 8.5.2), has Program several successful partnerships between CI and produced of a given companies to better understand the biodiversity In one in the earliest stages of development projects. area with forces example of this type of partnership, CI joined Alumina LLC and Alcan mining companies Alcoa World the diversity andInc. in 2005 to collect scientific data on coastal Guineastatus of species at a number of sites in considering placement of a the companies were where to In addition infrastructure. and related bauxite refinery ESIA, the survey also feeding information into the project made several important observations to advance scientific Notably for freshwater understanding of the region. crab, species, an Endangered was recorded for the first time since its original collection in was recorded 1947 (Cumberlidge 2006). 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An example of a species assessment summary and distribution map APPENDIX

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347 APPENDIX

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THE IUCN RED LIST OF THREATENED SPECIES ™

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