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Landscape Structure and Genetic Variation in Natural Zooplankton Populations

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Landscape Structure and Genetic Variation in Natural Zooplankton Populations LANDSCAPE STRUCTURE AND GENETIC VARIATION IN NATURAL ZOOPLANKTON POPULATIONS: CLADOCERANS IN A SET OF INTERCONNECTED PONDS ERIK MICHELS Promotoren: Prof. Dr. L. De Meester Prof. Dr. F. Ollevier Laboratorium voor Aquatische Ecologie Katholieke Universiteit Leuven Leden van de examencommissie: Prof. Dr. T. Backeljau Prof. Dr. L. Brendonck Prof. Dr. K. Schwenk Prof. Dr. F. Volckaert Het onderzoek voorgesteld in dit proefschrift werd uitgevoerd aan het Laboratorium voor Aquatische Ecologie (K.U.Leuven). Het onderzoek werd gesteund door een beurs verleend door het Instituut voor de bevordering van het Wetenschappelijk-Technologisch onderzoek in de industrie (I.W.T.). KATHOLIEKE UNIVERSITEIT LEUVEN FACULTEIT WETENSCHAPPEN DEPARTEMENT BIOLOGIE Laboratorium voor Aquatische Ecologie Landscape structure and genetic variation in natural zooplankton populations: cladocerans in a set of interconnected ponds Landschapsstructuur en genetische variatie in natuurlijke zoöplanktonpopulaties: cladoceren in een aantal onderling verbonden vijvers Promotoren: Prof. Dr. L. De Meester Proefschrift voorgedragen tot Prof. Dr. F. Ollevier behalen van de graad van Doctor in de Wetenschappen door Erik Michels 2001 Contents Dankwoord Chapter I General introduction and outline of the thesis: The use of a 1 landscape approach in population genetics Chapter II Zooplankton on the move: first results on the quantification of 39 dispersal of zooplankton in a set of interconnected ponds Chapter III Geographic and genetic distance among zooplankton populations 55 in a set of interconnected ponds: a plea for using GIS modelling of the effective geographic distance Chapter IV Microgeographic genetic structure of local Daphnia ambigua 75 populations connected by high dispersal rates Chapter V Genetic differentiation for ecologically relevant traits in a patchy 111 metapopulation of the cyclical parthenogen Daphnia ambigua Chapter VI Genetic differentiation in Polyphemus pediculus populations 137 General discussion 157 Summary 165 Samenvatting 171 Publications 177 Dankwoord Net als ik dacht dat ik er bijna was, besefte ik dat een dankwoord schrijven eigenlijk niet zo gemakkelijk is! Niet omdat ik het moeilijk heb om jullie te bedanken, wel omdat het gevaar groot is om in clichés te vervallen. Toch wil ik beginnen met iedereen te bedanken die in de voorbije jaren, ieder op zijn manier, er toe bijgedragen heeft dat dit boekje eindelijk voor u ligt. De voorbije jaren op het labo waren niet alleen wetenschappelijk zeer boeiend, maar gingen ook gepaard met de nodige toffe (en dikwijls memorable) momenten. Ik wil in de eerste plaats mijn promotoren bedanken die dit onderzoek in goede banen hebben geleid. Prof. Ollevier wil ik bedanken voor de gelegenheid die hij mij bood om in “het labo” steeds in de beste omstandigheden onderzoek te kunnen verrichten. Aan Luc De Meester heb ik ontzettend veel te danken. Luc’s wetenschappelijk inspiratie en enthousiasme werkten reeds tijdens het maken van mijn licentiaatsthesis aanstekelijk. Zonder “de hand van De Meester”, die ooit op een mooie zondagnamiddag in augustus de grote lijnen voor dit onderzoek op papier zette, zou dit doctoraat er dan ook nooit gekomen zijn. Luc zijn talloze commentaren en suggesties waren soms moeilijk leesbaar, maar brachten (eens ontcijferd) vaak een nieuw licht op dit onderzoek en werkten meestal zeer stimulerend. Ik ben Luc eveneens zeer dankbaar omdat hij mij de voorbije jaren ook de gelegenheid gaf om de wijde (wetenschappelijke) wereld te verkennen. Ik wil ook Luc Brendonck bedanken voor zijn niet aflatende interesse in dit onderzoek en zijn kritische maar constructieve commentaren op dit werk. Luc was reeds van bij het begin van dit onderzoek als “streng maar rechtvaardig IWT commissielid” bij dit onderzoek betrokken, maar ook nadien kon ik steeds bij hem terecht voor advies of gewoon een babbel. Bovendien apprecieer ik het ten zeerstse dat hij, na een gekneusde rib en enkele voltreffers, het nog steeds aandurft met mij te squashen. Hierbij wil ik ook de overige leden van de examencommissie: Prof. Filip Volckaert, Prof. Klaus Schwenk en Prof. Thierry Backeljau bedanken voor het grondig lezen en kritisch beoordelen van dit werk. Dit onderzoek zou onmogelijk geweest zijn zonder de gewaardeerde medewerking van de vzw Natuurreservaten (ondertussen Natuurpunt) en Willy Peumans, de conservator van De Maten. Ik besef maar al te goed wat een luxe het is geweest om te kunnen samenwerken met collega’s en studenten die ook goede vrienden geworden zijn. Dit doctoraat is voor een groot deel het resultaat van de wel bijzonder prettige samenwerking met Karl Cottenie die ik dan ook zeer dankbaar ben. Karl was mijn “partner in crime” tijdens het vele veldwerk in De Maten dat hierdoor bij momenten een avontuurlijk karakter kreeg en genoeg anekdotes opleverde voor menig straf verhaal. De bijdrage van Karl aan dit onderzoek beperkt zich echter niet tot het veldwerk of zijn statistisch advies (KISS* altijd en overal). Zijn inbreng in de uitwerking van de ideeën in dit werk zijn op minst aanzienlijk te noemen. Joost en Hanne, samen met Karl collega’s van het eerste uur, zorgden van in het begin mee voor de toffe werksfeer en zaten nooit verlegen om een wedstrijdje (s’ nachts op de fiets) of een weddenschap (ergens te velde). Ik wil zeker ook Lies (hoezo stress?) Neys en Koen (FT) De Gelas bedanken voor de zeer constructieve en plezante samenwerking tijdens hun thesis en ook erna. Lies wil ik speciaal bedanken voor haar hulp bij het maken van de figuur op de voorpagina van dit boekje. Katrien Uyttebroek zal ik mij altijd herinneren omdat ze zich tijdens haar thesis niet liet ontmoedigen door de moeilijkheden bij de kweek van D. ambigua. Verder wil ik Elke, Conny, Martine, An W., Ria, Tine, Els, Ellen, Steven, Frank, Joachim, Ronald, Eddy, Stef, Wouter, Jimmy, Joost R. Jochen, Gerald, Gregory, Jeroen en alle andere collega’s nog bedanken voor voor de toffe momenten aan de koffietafel en voor al die keren dat ze mij geholpen hebben bij allerhande kleine en minder kleine “doctoraatsperikelen”. Ik wil zeker ook niet nalaten al de vrijwilligers te bedanken die ooit hebben geholpen tijdens de jaarlijkse wonderbaarlijke visvangst in De Maten. Nicole Podoor en Wouter Rommens bedank ik voor hun hulp bij het analyseren van waterstalen uit De Maten en voor het bezorgen van gegevens over de waterplantenvegetatie in De Maten. Daarnaast zijn er natuurlijk nog al mijn vrienden die ik wil bedanken voor het bezorgen van de nodige (in)spanning en avontuur ter tijd en stond. Mijn (schoon)ouders, broer en zussen, bedank ik voor alles wat ze al die jaren voor mij gedaan hebben… Maar als er iemand is die ik niet genoeg kan bedanken is het Anne-Sophie. Zij heeft haar kersverse echtgenoot gedurende deze laatste drukke maanden vaak moeten missen en heeft alle “ups & downs” van dichtbij meegemaakt. Ik ben haar dan ook zeer dankbaar dat ze mij er steeds opnieuw van kon overtuigen dat het ook deze keer wel in orde zou komen. * Keep It Super Simple Introduction and outline Chapter I General introduction and outline of the thesis: The use of a landscape approach in population genetics Abstract This chapter presents an outline of the present work. In a short introduction, we point to the maintenance of genetic variation as a central aspect of conservation and evolutionary biology and underline the relevance of a landscape based metapopulation approach in population genetic research in zooplankton. Next, we briefly overview processes contributing to the genetic differentiation among zooplankton populations. In this respect we discuss patterns of local genetic differentiation as revealed by (quasi-)neutral markers and the study of ecological relevant traits and we focus at the difference between permanent and intermittent populations of cyclical parthenogens. In addition, we point at the extensive genetic polymorphism for predator induced defence mechanisms. As metapopulation biology provides a useful theoretical framework to study genetic differentiation in a set of zooplankton populations linked by dispersal, we introduce the basic concepts of metapopulation biology and highlight important recent developments in this field, i.e. the synthesis of metapopulation studies and landscape ecology. From this overview, we argue that a landscape based approach may offer an attractive framework for population genetic studies. At the end of this chapter we provide some historical and ecological background information to the study area, the pond complex of 1 Chapter I the nature reserve De Maten and present the species characteristics of the two model species, Daphnia ambigua (Scourfield, 1947) and Polyphemus pediculus (Müller, 1785). Introduction Genetic variation is a conditio sine qua non for evolution, since it enables natural populations to adapt genetically to the constantly changing environmental conditions. During the last three decades, the maintenance of genetic diversity in natural populations has been a central issue in evolutionairy biology and a major concern in conservation biology (Soulé & Simberloff, 1986; Lande, 1988; Loeschke et al., 1994; Frankham, 1995; Avise & Hamrick, 1996; Spellerberg, 1996; Meffe & Caroll, 1997). Since the early studies by Lewontin in 1974, genetic studies using allozyme markers and more recently polymerase chain reaction enhanced DNA techniques have revealed a substantial genetic diversity in most natural populations. Several hypotheses have been proposed to explain the maintenance of genetic diversity in populations.
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