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Ecological and Genetic Effects of Urbanization and Fragmentation on Carabid Beetles from Belgian Beech Forests

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Ecological and Genetic Effects of Urbanization and Fragmentation on Carabid Beetles from Belgian Beech Forests Universiteit Gent Faculteit Wetenschappen Academiejaar 2011-2012 Ecological and genetic effects of urbanization and fragmentation on carabid beetles from Belgian beech forests Ecologische en genetische effecten van urbanisatie en fragmentatie op loopkevers in Belgische beukenbossen door Eva Gaublomme Thesis submitted in fulfillment of the requirements for the degree of Doctor (Ph.D.) in Sciences Proefschrift voorgedragen tot het behalen van de graad van Doctor in de Wetenschappen Promotor: Prof. Dr. Johan Mertens (Ghent University) Co-promotor: Prof. Frederik Hendrickx (Ghent University – RBINSc) Co-promotor: Dr. Konjev Desender† (RBINSc) Reading committee: Prof. Carl Vangestel (Ghent University – UC Davis, Califronia, USA) Dr. Claudia Drees (Leuphana Universität Lüneburg, Germany) Dr. Wouter Dekoninck (RBINSc) Examination committee: Prof. Johan Mertens (Ghent University, Promotor) Prof. Frederik Hendrickx (Ghent University - RBINSc, Co-promotor) Prof. Luc Lens (Ghent University, chairman) Prof. Carl Vangestel (Ghent University – UC Davis, Califronia, USA) Dr. Claudia Drees (Leuphana Universität Lüneburg, Germany) Dr. Wouter Dekoninck (RBINSc) ISBN number University Press, Zelzate Please refer to this work as: Gaublomme, E. (2012). Ecological and genetic effects of urbanization and fragmentation on carabid beetles from Belgian beech forests. PhD thesis, Ghent University. In memoriam of Konjev Table of contents CHAPTER 1 INTRODUCTION AND THESIS OUTLINE 1 CHAPTER 2 THE EFFECTS OF FOREST PATCH SIZE AND MATRIX TYPE ON CHANGES IN CARABID BEETLE ASSEMBLAGES 27 CHAPTER 3 LOCAL EXTINCTION PROCESSES RATHER THAN EDGE EFFECTS AFFECT GROUND BEETLES ASSEMBLAGES IN FRAGMENTED OLD BEECH FORESTS . 57 CHAPTER 4 ISOLATION AND CHARACTERIZATION OF MICROSATELLITE LOCI IN THE GROUND BEETLE CARABUS PROBLEMATICUS (C OLEOPTERA , CARABIDAE ) 79 CHAPTER 5 NON -DESTRUCTIVE SAMPLING FOR GENETIC STUDIES ON CARABUS AURONITENS AND CARABUS PROBLEMATICUS : A STUDY BASED ON ALLOZYMES AND MICROSATELLITES 89 CHAPTER 6 LOSS OF GENETIC DIVERSITY AND INCREASED GENETIC STRUCTURE IN RESPONSE TO FOREST AREA REDUCTION IN THE FLIGHTLESS CARABID BEETLE CARABUS PROBLEMATICUS (C OLEOPTERA : CARABIDAE ). 101 CHAPTER 7 IMPACT OF FRAGMENTATION ON THE GENETIC STRUCTURE OF THE FOREST GENERALIST CARABUS VIOLACEUS , WITH COMPARISON TO THE FOREST SPECIALIST CARABUS PROBLEMATICUS (C OLEOPTERA : CARABIDAE ) 129 CHAPTER 8 GENERAL DISCUSSION 157 SUMMARY 173 SAMENVATTING 177 Dankwoord Dankwoord In de allereerste plaats gaat mijn grote dankbaarheid uit naar Konjev Desender. Ook al is hij er niet meer, hij is me steeds blijven volgen via het gedenkkaartje dat bovenaan mijn bureau staat. Hij heeft de weg voor mij geopend en daarvoor ben ik hem zeer dankbaar. Vervolgens gaat mijn dank uit naar mijn promotor, Johan Mertens die me jaar na jaar is blijven steunen, en het waren er vele. Van onschatbare waarde was de hulp die ik ondervond van Frederik. Stiekem noemden we hem de blonde halfgod. Je slaagde er telkens opnieuw in om me te motiveren en daarvoor Frederik kan ik jou nooit genoeg bedanken. Zonder jou was dit doctoraat nooit afgewerkt. Om op hetzelfde elan verder te gaan, wil ik ook de aartsengel Thierry bedanken die me op de onmogelijkste uren papers doorstuurde, al dan niet op aanvraag. Maar zeker ook voor de morele steun die ik van jou kreeg. Je staat altijd klaar voor iedereen, dag en nacht. Bedankt Thierry. Verder wil ik ook het voltallig departement Entomologie geleid door Patrick Grootaert bedanken. En hierbij ook reeds talloze collega’s die nu op andere terreinen actief zijn. Ik wil echter in het bijzonder Alain bedanken. Alain, merci pour tous les triages, les jours de terrains et les électrophorèses. On en a fait de nombreux. Avec toi c’était toujours très agréable, sauf dans le ‘Strontbos’. Ook iedereen uit het labo en gedurende die hele periode zijn er heel wat de revue gepasseerd. Graag wil ik Karin, Charlotte, Hilde V, Hilde D, Hilde E, Guy, Karine, Patrick, Tine, Dries, Isa, Zohra, Lynn, Gontran, Zoltan, Massi, Vanya en Charlotte bedanken en verder nog een hele rits buitenlanders die voor variabele tijd het labo kwamen opvrolijken. Aan de ijsgevechten kwam helaas een einde, want de machine heeft de geest gegeven. Verder wil ik in het bijzonder Veerle, Domir, Karine en Karin bedanken. Zij hebben mij van begin tot einde gesteund en die steun was van onschatbare waarde. Ook de steun van Dankwoord Charlotte en Hilde E tijdens de laatste jaren deed enorm goed en daarom een heel wel gemeende dikke merci voor jullie jarenlange geduld. Ook op het terrein werd ik niet enkel bijgestaan door Alain, maar ook door ons vake die om de veertien dagen gedurende 38 staalnames trouw met mij op pad ging om de bossen in Jette onveilig te maken of eerder veilig te maken voor mij. Ook ons moeke, tante Marianne, tante Monique en tante Christelle zijn met mij door de bossen getrokken om de bodemvallen te ledigen. Dank jullie wel voor dit leuke gezelschap tijden de vele staalnames die op mijn agenda stonden. Ook Wouter F wil ik graag bedanken voor de hulp bij het extraheren en pcr’en van de talloze specimens. Binnenkort verdedig jij ook en als de baby het toelaat kom ik zeker luisteren. Ook in de Ledeganck ben ik heel veel mensen dank verschuldigd. Het was daar soms een zotte bende, zeker in bureau de Coolerds met Debbie, Kevin, Boris, Martijn en Sudhi. Ook de koffiepauzes, al dan niet met koekjes, waren een leuke afwisseling. Bedankt iedereen voor de fijne periode in Gent. Tim, jij verdient ook een speciaal woordje in dit hoofdstuk. Talloze keren heb jij voor mij Gis kaartjes gemaakt die telkens na commentaren van revisoren moesten worden aangepast. Bedankt voor al jouw geduld en je hulp. Ook op het INBO wil ik in de eerste plaats Bruno bedanken alsook Mathieu en ‘de twee Koens’. Met jullie op terrein gaan was zeer leuk, op de vroege ochtenduren na voor het nemen van de fish eye foto’s. Verder ook nog vele vriendjes die zorgden voor de nodige ontspanning en gelukkig niet teveel vroegen naar de vooruitgang, want soms ging het tergend traag. Inge, Tine, Katinka en Gwendolyn, volgend seizoen doe ik hopelijk terug mee met de badminton competitie. Ook de meisjes Kaatje, Ann, Leen, Babs, merci voor de ontspannen etentjes. En verder ook Jan, Eva, Nic, de bende van ‘den donderdagavond’ en zoveel anderen. Bedankt voor de ontspanning tussendoor. Dankwoord Mijn schoonouders wil ik ook heel graag bedanken Vooral voor het opvangen van de kindjes tijdens de laatste, drukke maanden vóór het indienen. Het was echt super dat ik af en toe eens heel hard kan doordoen zonder achtergrondmuziek. Ook broertje en schoonzusje Ellen, bedankt voor de steun en Ellen bedankt voor de opsmuk van de overbelichte foto’s die ik nam van ons vake zijn schilderijen. Maar van onschatbare waarde, zijn mijn ouders. Toen ze vorig jaar ons gezinnetje van vier opnamen in hun huis zullen ze wel niet gedacht hebben dat het zo druk zou worden; niet alleen waren er de verbouwingen, daarbij kwam nog eens het niet altijd even goede humeur van dochterlief wegens hormonenwissels door de bollende buik en dan nog eens de zware periode van “het afwerken”. Indien we niet bij jullie hadden gewoond, was het mij zeker nooit gelukt. Jullie namen bijna het volledige huishouden op jullie schouders, gecombineerd met meehelpen op de werf, ongelooflijk veel voor de kindjes zorgen en al de rest. Het is dan ook nooit mogelijk om jullie voldoende te bedanken voor alle kansen die jullie mij hebben geboden. Toch wil ik een poging ondernemen en vanuit het diepste van mijn hart zeg ik jullie: Heel erg bedankt! Jullie zijn fantastisch. En last but not least mijn drie ventjes. De twee kleinsten hielpen het niet echt altijd vooruit, maar vrolijkten me wel op als het eens wat minder goed ging. Hopelijk komen er nu veel roze dagen en hoeven we geen kruisjes meer te zetten tot wanneer mama eindelijk eens wat minder moet werken. Jakob stuurde me steevast naar mijn bureau als ik eens even in de zetel zat en Elias hield zich samen met broer trouw aan de wekker. Als hij afloopt mogen we mama gaan storen. “Joepie”. Nu zal ik eindelijk wat meer tijd hebben en gaan we terug als gezin op pad in plaats van alleenstaande papa Bart. Kleine zus in mijn buik schopte me wel wakker, moest ik ooit dreigen in slaap te vallen. Mijn allergrootste dank gaat echter uit naar de grootste van de drie. Mijn allerliefste poepie. Jij bent me altijd blijven steunen, al geloofde je er zelf misschien niet altijd meer in. Dankzij jou is het ‘eindelijk’ tot een goed einde gebracht en nu gaan we genieten genieten en nog eens genieten. Binnenkort met een nieuw creatuurtje erbij. Hopelijk wordt het geen ADHD-erke. Ik zie jou ontzettend graag. Bedankt voor alles, mijn super grote schat. Painting of fish eye photo for light measurements in the forest by Luc Gaublomme © Eva Gaublomme ___________________________________________________________________________ Chapter 1 INTRODUCTION AND THESIS OUTLINE ___________________________________________________________________________ Introduction and thesis outline Chapter 1 Introduction and thesis outline ___________________________________________________________________________ 1.1 Area and isolation effects as a consequence of habitat fragmentation Landscape modification for agriculture and urbanization puts severe stress on natural ecosystems because of increasing habitat loss and deteriorating habitat quality. Forest fragmentation is an important process contributing to the present-day concern over the loss of biodiversity and rates of species extinction (Meffe & Carroll 1997, Fahrig 2003, Ewers & Didham 2006). It results in a landscape composed of habitat fragments (e.g. forest) with something else (the matrix) between the fragments. This results in a decrease of population size and increased isolation of habitat patches (MacArthur & Wilson 1967). Besides the direct reduction in population size, small populations become more prone to extinction as stochastic variation may result in a higher probability of extinction in small compared to large populations. The assumed processes likely to be involved in species extinctions as consequences of habitat fragmentation have been conceptualized in the so-called extinction vortex (Frankham et al.
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