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Distribution of Spiders in Coastal Grey Dunes

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Distribution of Spiders in Coastal Grey Dunes kaft_def 7/8/04 11:22 AM Pagina 1 SPATIAL PATTERNS AND EVOLUTIONARY D ISTRIBUTION OF SPIDERS IN COASTAL GREY DUNES Distribution of spiders in coastal grey dunes SPATIAL PATTERNS AND EVOLUTIONARY- ECOLOGICAL IMPORTANCE OF DISPERSAL - ECOLOGICAL IMPORTANCE OF DISPERSAL Dries Bonte Dispersal is crucial in structuring species distribution, population structure and species ranges at large geographical scales or within local patchily distributed populations. The knowledge of dispersal evolution, motivation, its effect on metapopulation dynamics and species distribution at multiple scales is poorly understood and many questions remain unsolved or require empirical verification. In this thesis we contribute to the knowledge of dispersal, by studying both ecological and evolutionary aspects of spider dispersal in fragmented grey dunes. Studies were performed at the individual, population and assemblage level and indicate that behavioural traits narrowly linked to dispersal, con- siderably show [adaptive] variation in function of habitat quality and geometry. Dispersal also determines spider distribution patterns and metapopulation dynamics. Consequently, our results stress the need to integrate knowledge on behavioural ecology within the study of ecological landscapes. / Promotor: Prof. Dr. Eckhart Kuijken [Ghent University & Institute of Nature Dries Bonte Conservation] Co-promotor: Prf. Dr. Jean-Pierre Maelfait [Ghent University & Institute of Nature Conservation] and Prof. Dr. Luc lens [Ghent University] Date of public defence: 6 February 2004 [Ghent University] Universiteit Gent Faculteit Wetenschappen Academiejaar 2003-2004 Distribution of spiders in coastal grey dunes: spatial patterns and evolutionary-ecological importance of dispersal Verspreiding van spinnen in grijze kustduinen: ruimtelijke patronen en evolutionair-ecologisch belang van dispersie door Dries Bonte Thesis submitted in fulfilment of the requirements for the degree of Doctor [Ph.D.] in Sciences Proefschrift voorgedragen tot het bekomen van de graad van Doctor in de Wetenschappen Promotor: Prof. Dr. E. Kuijken Co-promotoren: Prof. Dr. J.-P. Maelfait & Prof. Dr. L. Lens D ANKWOORD Zowel genen als omgeving bepalen de levensloop van mens en dier. Zonder dat ik die eerste oneer wil aandoen, heeft de omgeving in mijn geval een bepalende invloed gehad op mijn interesses en de thematiek van dit proef- schrift, aangezien ik opgroeide te midden van de laatste restanten kustdui- nen. Daarom ben ik bijzonder trots om dit werk te kunnen voorleggen als wat ooit met vogels-kijken in de duinen begon. Hoewel alleen mijn naam boven aan dit proefschrift staat, kon het enkel tot stand komen met de niet te onderschatten steun en medewerking van collega’s, vrienden en familie. In de eerste plaats ben ik veel dank verschuldigd aan mijn promotor Eckhart Kuijken, die mij van het begin af gestimuleerd heeft om een type onderzoek te verrichten waarin ganzen geen enkele rol spelen. Nog belangrijker was de voortreffelijke begeleiding en onbaatzuchtige steun van Jean-Pierre Maelfait. Zonder de vele [gemoedelijke] discussies, vaak tussen pot en pint, en zijn kritische opvolging zou ik dit proefschrift nooit tot een goed einde hebben kunnen brengen. Luc Lens ben ik heel dankbaar voor het aanleren van SAS en voor zijn constructieve medewerking aan de afwerking van enkele manu- scripten, wat de kwaliteit ervan op een hoger niveau gebracht heeft. Jean- Pierre en Luc, duizendmaal dank, vanuit de grond van mijn hart! I am also grateful to the other members of the examination-committee for their valuable comments and suggestions on the preliminary version of this thesis. Alle collega’s van het negende wil ik uitdrukkelijk danken voor de fijne werk- sfeer tijdens en na de ‘werk’uren, waardoor afzakken naar het labo steeds een plezier was [en is]. Frederik, bedankt voor de vele discussies en de, mis- schien onbewuste, stimulans om steeds beter onderzoek te leveren. Verder wil ik ook Danny en Viki speciaal vermelden voor de gezellige babbels aan de koffietafel, de hulp bij ‘de kweek’ en jullie interesse in mijn onderzoek. Domir, Shirley en Wouter, jammer dat jullie niet op het negende konden blijven, want dankzij jullie was ambiance steeds verzekerd. Etienne, bedankt voor het in stand houden van de collembola-kweek! Tenslotte wil ik ook Jeroen, Lynda en Angelica noemen, die me hebben geholpen bij kleine en grote PC-, print-, scan- en SAP-problemen. 2 Tijdens de begeleiding van de vele scriptie- en maandwerkstudenten kreeg ik niet enkel de kans om op een boeiende manier mijn interesse in spinnen en ecologie door te geven, maar heb ik tevens heel wat gegevens kunnen verza- melen die van essentieel belang waren voor de afwerking van dit proef- schrift. Dank aan de ‘determineerders’ Tom, Peggy, Isabelle en Liesbeth en de ‘ballooners’ Isra, Nele en Jeroen voor hun onmisbare bijdragen! De practica, excursies en stages waren voor mij een leerrijke en aangename afwisseling tijdens mijn assistentschap. Hendrik, Hara, Eric, Sandra, Wim, Bart, Wouter, Ruben en Sam: bedankt voor de leuke samenwerking en ont- spannende momenten. Ook mijn welgemeende dank aan de andere colle- ga’s van de vakgroep, het I.N. en het K.B.I.N., met wie ik tijdens allerlei onderzoeksprojecten in contact kwam: Léon, bedankt voor je spinnengege- vens; Sam, we moeten dringend nog eens vergaderen met Bowmore, en Patrick, jouw enthousiasme zal ik niet snel vergeten! Uiteraard ben ik ook mijn ouders ontzettend dankbaar voor hun stimulerend enthousiasme en hun niet-aflatende appreciatie voor mijn belangstelling in wat er zoal groeit en kruipt in duin en achtertuin. Ook de rest van de familie en schoonfamilie wens ik bij deze uitdrukkelijk te vermelden vanwege hun begrip [ ‘Moe je gie nu were kobben goan vangen in de dunen ?’] en logistieke steun tijdens en na het veldwerk [catering is nu eenmaal heel belangrijk]. Roxane en Kasper, jullie komen als laatste aan bod. Rox, ik moet toegeven, samenleven met een door spinnen geintrigeerde veldbioloog is waarschijnlijk niet altijd even gemakkelijk. Je hebt dat goed gedaan! Kasper, tenslotte, jij ‘verplichtte’ mij om de nodige ontspanning te nemen, hoewel dat daarom ’s nachts echt niet hoefde hoor... Bedankt voor jullie steun, liefde en het goede gevoel dat jullie me gaven !!! Dries Bonte, 26 januari 2004 HOOFDSTUK / 3 Andere leden van de lees- en examencommissie: Dr. J. Bell [Horticulture International, Wellesbourne, UK] Prof. Dr. M. Hoffmann [Universiteit Gent] Prof. Dr. J. Mertens [Universiteit Gent] Dr. H. Van Dyck [Universiteit Antwerpen] Prof. Dr. M. Vincx [Universiteit Gent, Voorzitter] 4 T ABLE OF CONTENTS Dankwoord 2 Chapter I Thesis outline and introduction 6 ASSEMBLAGE STRUCTURE Chapter II.1 An introduction to spider assemblages of coastaldunes 17 Chapter II.2 Spider assemblage structure and stability in a heterogeneous coastal dune system 22 Chapter II.3 Regional and local variation of spider assemblages [Araneae] from coastal grey dunes along the North Sea 40 Chapter II.4 Spider assemblages from coastal dunes: general discussion 60 EVOLUTIONARY ECOLOGY OF DISPERSAL Chapter III.1 Evolutionary and ecological aspects of dispersal, with emphasis on spider dispersal behaviour 68 Chapter III.2 Low propensity for aerial dispersal in specialist spiders from fragmented landscapes 96 Chapter III.3 Effects of aerial dispersal, habitat specialisation, and landscape structure on spider distribution across fragmented grey dunes 109 Chapter III.4 Environmental and genetic background of tiptoe-initiating behaviour in the dwarfspider Erigone atra 123 Chapter III.5 Patch quality and connectivity influence spatial dynamics in a dune wolfspider 137 Chapter III.6 Evolution of ballooning behaviour in a wolf spider inhabiting different landscapes 154 Chapter III.7 Lack of homeward orientation and increased mobility result in high emigration rates from low-quality fragments in a dune wolf spider 175 Chapter III.8 Spider dispersal in fragmented coastal grey dunes: general discussion 192 Chapter IV.1 Considerations for nature conservation and future research 204 References 214 Samenvatting 235 Curriculum Vitae 244 TABLE OF CONTENTS / 5 I.1T HESIS OUTLINE AND INTRODUCTION foto: dries bonte 6 Objective and outline of the thesis This Ph.D. study contributes to the ‘Nature management monitoring pro- gramme of the Belgian coastal dunes’ [1996-2002]. This programme is based on a multidisciplinary, integrated approach. The three main research enti- ties participating in this programme are the Laboratory of Soil Science and Terrestrial Ecology Unit of Ghent University and the Institute of Nature Conservation [Section Landscape Ecology and Nature Management]. The motherproject was funded by the Ministry of the Flemish Community, AMI- NAL, department Nature and initially [1996-1999] cofinanced by the LIFE- project ‘Integrated Coastal Conservation Initiative [ICCI]’. The primary aim of the monitoring programme is to evaluate the results of the applied nature management measures, especially grazing by domestic livestock, in the coastal dunes, but additional research projects were started up in the Terrestrial Ecology Unit of Ghent University as an extension of this descrip- tive monitoring. My research project, funded by Ghent University, did not aim to evaluate management actions senso stricto but had the objective to document and predict variation in spider distribution in function of a changing landscape, where grazing is a tool to interconnect and enlarge the remaining grey dune fragments. Dispersal behaviour, a key
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