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Regional Diversity, Ecology and Palaeoecology of Aquatic Invertebrate Communities in East African Lakes

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Regional Diversity, Ecology and Palaeoecology of Aquatic Invertebrate Communities in East African Lakes Academic year 2009-2010 Thesis submitted in partial fulfillment of the requirements for the degree of Doctor in Science: Biology Regional diversity, ecology and palaeoecology of aquatic invertebrate communities in East African lakes Regionale diversiteit, ecologie en paleo-ecologie van aquatische invertebraten- gemeenschappen in Oost-Afrikaanse meren BOB RUMES Promotor: Prof. Dr. D. Verschuren Co-promotor: Dr. H. Eggermont Faculty of Sciences, Biology Department Research group Limnology Members of the examination committee Members of the reading committee Promotor: Prof. Dr. D. Verschuren (Universiteit Gent) Co-promotor: Dr. H. Eggermont (Universiteit Gent) Prof. Dr. K. Martens (Koninklijk Belgisch Instituut voor Natuurwetenschappen) Prof. Em. Dr. H. Dumont (Universiteit Gent) Dr. O. Heiri (Universiteit Utrecht) Other members of the examination committee Prof. Dr. D. Adriaens (Universiteit Gent) (chairman) Prof. Dr. W. Vyverman (Universiteit Gent) Public thesis defence Thursday, March 11th, 2010 at 4 p.m. Ghent University, campus Ledeganck, Auditorium 4, K.L. Ledeganckstraat 35, 9000 Gent This thesis was realised within the framework of CLANIMAE (Climatic and Anthropogenic Impacts on African Ecosystems) funded by the Belgian Science Policy (BelSPO). Dankwoord Dit doktoraat zou er niet zijn zonder de hulp van een hele hoop mensen die (hoofdzakelijk) vrijwillig hun tijd in mij hebben geïnvesteerd. Een kort woordje van dank is dan ook niet ongepast. In de eerste plaats wil ik Dirk Verschuren bedanken. Dirk, bedankt voor het vertrouwen en het geduld. Het is niet altijd vlot verlopen en soms leek het alsof elke keer dat we samenzaten de structuur van mijn doktoraat veranderde, maar in het uiteindelijke resultaat kunnen we ons beiden terugvinden. De hulp die ik heb gekregen van Hilde Eggermont, mijn (vrijwel) immer blijgezinde en enthousiaste co-promotor, stel ik ook bijzonder op prijs. Ik kreeg van jou snelle en goede feedback, oplossingen voor problemen en je had altijd een kop warme koffie beschikbaar. Wat meer kan je wensen een co- promotor? Dat ik geen koffie lust, is jouw schuld niet. Ik bedank bij deze ook Prof. Henri Dumont, die me al sinds 2000 een plaats heeft gegeven op zijn labo en dit ondanks het feit dat ik eerst enkel op zoek was naar transeksuele pissebedden. Toen ik me voor mijn doktoraat ben beginnen interesseren in de watervlooien hebben hij en Kay Van Damme me goed vooruit geholpen. Prof. Koen Martens van het KBIN bedank ik o.a. voor de continue steun in mijn werk op de ostracoden. De platen van de ostracoden zijn ook een verdienste van Julien Cillis wiens vaardigheid de temperamentvolle scanning electronen microscoop van het KBIN keer op keer terug aan de praat kreeg. Ook de andere leden van de examenjury, Oliver Heiri, Wim Vyverman en Dominique Adriaens bedank ik voor hun feedback op de draft versie van deze thesis en hun aanwezigheid op de voor- en publieke verdediging. Daarnaast dank ik Lei Chou (ULB), Pierre-Denis Plisnier en Harlod Hughes (KMMA) voor de analyses van waterchemie, Ilse Bessems voor de sedimentologische data, Dirk Van Damme voor hulp bij de identificatie van de Oegandese mollusken, Joachim Mergeay voor de staalnames in Kenia, Julie Lebrun voor informatie over de Oegandese waterplanten, het laboratorium voor houttechnologie voor het gebruik van hun microscoop met camera, Angelica Alcantara voor all- round assistentie in het labo, Vanessa Gelorini om tilia helpdesk te spelen, Thijs Van der Meeren om de vele obscure referenties van tropische ostracoden te zoeken en Générose Kabihogo voor het geduld en het doorzettingsvermogen waarmee ze duizenden invertebratenresten heeft uitgepikt. Voor de sfeer op het labo dank ik zowel radiolabo - met DJ’s Hilde, Kay, Deo, Ilse, Supiyanit, Maureen en Angelica – als de jongere garde van Labo Limno met Générose, Saber, Els, Christine C., Leen, Thijs, Vanessa en Joeri. Ik heb ook veel goede herinneringen aan de mensen van de TEREC die ik af en toe van kantoormateriaal/beamer heb voorzien (plaatsvervangende Angelica) en die ik verder voornamelijk ontmoette op feestjes, recepties en bij de lunch. Zonder veldwerk zou er verderop in de thesis niet veel staan. Ik bedank het Leopold III-Fonds voor natuuronderzoek en natuurbehoud, het Fonds Wetenschappelijk Onderzoek Vlaanderen (FWO) en de Vlaamse Interuniversitaire Raad (VLIR) voor hun financële steun. I’m also grateful to Pierre-Denis Plisnier, Maureen Fagot, Vanessa Gelorini, Julie Lebrun, Johnson Bwembale, Julius Lejju and Imke De Windt for their assistance in the field. I would like to thank the Uganda National Council for Science and Technology and the Uganda Wildlife Authority for granting us the necessary permits to do this research. I’m especially gratefull to the ugandans I met while conducting the work. Most of you were very kind, interested in the work and often offered assistance or valuable information. If anyone is to benefit from this study it should be you. Bij deze wil ik ook de collega’s van de BMM bedanken die me de laatste maanden regelmatig hebben moeten missen omdat ik ‘nog even iets moest afwerken’. In het bijzonder wil ik Thierry Jacques bedanken die me altijd gesteund heeft in de beslissing om mijn doktoraat af te werken. Duiken, lezen, gezelschapsspelen, uitgebreid tafelen en regelmatig een pintje gaan pakken (al dan niet na een zware BDW-travelling excursie) – en dit alles in een bont gezelschap van oud-biologen, ere- biologen, Gudders, Mummies, Trollen, accesoires en ander staatsgevaarlijk gespuis - hebben ervoor gezorgd dat de boog niet altijd gespannen stond. Bedankt! Tot slot bedank ik ook mijn familie en ouders die me gedurende vele jaren gesteund hebben en nu ongetwijfeld blij zijn met het feit dat ik eindelijk ‘afgestudeerd’ ben. De kans is klein dat ik bij de volgende familiefeestjes in Afrika zit. Imke, de beste manier om jou te bedanken ligt voor de hand. Bedankt allemaal! Table of contents i Table of contents Table of contents i Samenvatting iii Summary vii Introduction 1 Chapter 1 Environmental regulation of the distribution and faunal richness of Cladocera in western Uganda crater lakes 13 Chapter 2 Environmental regulation of ostracod (Crustacea: Ostracoda) communities in western Uganda crater lakes. 39 Chapter 3 Ostracod (Ostracoda, Crustacea) communities in shallow Kenyan lakes and ponds 65 Chapter 4 Ecological and taphonomic controls on the composition of fossil aquatic invertebrate assemblages across a wide environmental gradient of East African lakes 87 Chapter 5 A four thousand year history of crater lakes in western Uganda: response of the aquatic invertebrate fauna to climatic and anthropogenic impacts 125 Chapter 6 Climatic and hydrological instability in semi-arid equatorial East Africa during the late Glacial to Holocene transition: A multi-proxy reconstruction of aquatic ecosystem response in northern Tanzania. 167 Chapter 7 Aquatic community response in a groundwater-fed desert lake to Holocene desiccation of the Sahara. 195 General Discussion 229 General Conclusion 247 Appendix 255 ii Samenvatting iii Samenvatting Gezien de wereldwijde toename van het gebruik van land door de mens is het een belangrijke uitdaging voor aquatische ecologen om te bepalen in welke mate deze menselijke activiteiten een invloed hebben op de structuur en de functie van aquatisch ecosystemen. In dit opzicht moet men ook het herstellingsvermogen van de aangetaste waters inschatten (Sutherland et al, 2002). In Afrika wordt verwacht dat klimaatveranderingen en de klimaatschommelingen extra druk zullen plaatsen op zowel de beschikbaarheid van en de toegang tot (zoet) water als op de vraag ernaar (Boko et al., 2007). Een grondige kennis over de respons van zoetwaterecosystemen op vroegere variaties in klimaat en menselijke invloed is essentieel om een beter inzicht te verwerven in de veerkracht van deze ecosystemen. Dit laat ons toe om een inschatting te kunnen maken van hun veerkracht ten opzichte van de voorspelde klimaatverandering en de verwachte toename van intensief landgebruik. Hetgeen op zijn beurt noodzakelijk is voor het opstellen van een beheersstrategie die het behoud van de functies en de diensten van het ecosysteem op lange termijn weet te handhaven. Deze doctoraatsstudie wil de respons en het herstelvermogen van Oost-Afrikaanse meren op veranderingen in het milieu nagaan. De nadruk wordt hierbij gelegd op de gevoeligheid van lokale aquatische gemeenschappen voor abiotische veranderingen in hun omgeving alsook op de snelheid en de wijze van het herstel na een periode van verstoring. In dit onderzoek werden de natuurlijke klimaatsgedreven veranderingen in het aquatisch milieu (variatie in habitat gerelateerd aan veranderingen in meerniveau en saliniteit) onderzocht evenals de veranderingen door menselijke verstoring van het meer en het omringend terrestrisch ecosysteem. Methodologisch gezien hebben we ons hierbij geconcentreerd op het potentieel van een aantal tot nog toe weinig gebruikte zoölogische groepen uit het sedimentarchief van Afrikaanse meren als indicatoren voor omgevingsveranderingen. In een eerste onderzoeksluik werden de hedendaagse ongewervelde zoetwatergemeenschappen uit West-Oeganda (hoofdstuk I, II en IV) en Zuid-Kenia (hoofdstuk III) bestudeerd. Op deze manier werd nagegaan welke omgevingsparameters de soortensamenstelling bepaalt. De verspreiding van Cladocera (watervlooien; Crustacea: Branchiopoda) in Oegandese meren werd het sterkst beïnvloed door de beschikbaarheid van nutriënten (totaal fosforgehalte, TP), de aanwezigheid en diversiteit van waterplanten,
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