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Het Lot Van Endozoöchoor Verbreide Zaden The fate of endozoochorously dispersed seeds Het lot van endozoöchoor verbreide zaden Tanja Milotić Promotor: Prof. Dr. Maurice Hoffmann Ghent University Faculty of Sciences Academic Year 2016-2017 Submitted to the faculty of Sciences of Ghent University, in fulfilment of the requirements for the degree of Doctor (PhD) in Sciences Members of the examination committee Prof. Dr. Maurice Hoffmann (Ghent University/INBO, Belgium): promotor Prof. Dr. Dries Bonte (Ghent University, Belgium): chair Prof. Dr. Annemieke Verbeken (Ghent University, Belgium): secretary Dr. Bram D'hondt (Ghent University/ANB, Belgium) Dr. Amy Eycott (University of Bergen, Norway) Prof. Dr. Robin Pakeman (James Hutton Institute, UK) Prof. Dr. Kris Verheyen (Ghent University, Belgium) Date of public defence: February, 24th 2017 For citation of published work reprinted in this thesis, please cite the original publication as mentioned at the first page of each chapter. Please cite this thesis as: Milotić, T. (2017) The fate of endozoochorously dispersed seeds. PhD dissertation. Ghent University, Ghent, Belgium. pp 360. The research in this thesis was conducted at the Terrestrial Ecology Unit (TEREC), Biology Department, Faculty of Sciences, Ghent University K.L. Ledeganckstraat 35, 9000 Ghent, Belgium Pictures taken by Tanja Milotić unless stated otherwise Acknowledgments Na zes jaar zweten en zwoegen ligt het er eindelijk: mijn doctoraat! Ontelbaar vaak heb ik op vragen moeten antwoorden met "neen, mijn doctoraat is nog niet af, maar ik heb nog massa's tijd" of "neen, ik ben helemaal geen post-doc maar gewoon een overjaarse (?) doctoraatsstudent" (jaja, ik ben er iets rijper aan begonnen dan gebruikelijk). Ondertussen leek die 'massa tijd' exponentieel af te nemen terwijl het met het aantal ideeën over extra toe te voegen hoofdstukken eerder de omgekeerde richting uitging. Zonder de hulp van een heleboel mensen had dit doctoraat er nooit gelegen, dus bij deze een poging om jullie allemaal te bedanken… Eerst en vooral zou ik mijn promoter, Maurice Hoffmann, willen bedanken om me de kans te geven om dit doctoraat te schrijven en de vrijheid die ik hierbij gekregen heb. Ik kende je al voordat ik aan dit avontuur begon uit een vorig leven bij het INBO, en gezien je je handen wel vol hebt aan het INBO heb je me van in het begin al duidelijk proberen te maken dat zelfstandig werken een absolute must zou zijn tijdens dit doctoraat. Anderzijds denk ik niet dat elke doctoraatstudent kan zeggen bloempotten gevuld te hebben met zijn/haar promotor, of in de kustreservaten op zoek is geweest naar geschikte plekjes voor veldexperimenten. Dus dat afwezig zijn viel al bij al bijzonder goed mee want het is vooral gebleken dat ik steeds op je kon rekenen wanneer ik je nodig had. Je bracht me in contact met de juiste mensen, inspireerde me, ronselde bachelor- en master-studenten en las bijzonder veel teksten na, zeker naarmate de deadline naderde. Zonder jouw hulp en enthousiasme had dit resultaat er nu niet gelegen… I also would like to thank the members of the examination committee for accepting this 'Christmas gift', and providing ideas and corrections which definitely improved this thesis. I realise that my PhD is rather voluminous while your agenda is probably overloaded. Still you managed to notice the forgotten parts, wrong subtitles and suggest corrections to improve the text. I really appreciate your effort! Verder zou ik graag alle collega's van de TEREC en LIMNO willen bedanken voor de 6 fijne jaren die ik op de universiteit mocht doorbrengen! Gezien mijn vergeetachtig karakter durf ik het niet aan om al jullie namen op te i lijsten, maar bedankt om voor de aangename sfeer op de unief en ver daarbuiten (bijv. tijdens de stages). Mijn bureaugenootjes Valérie Lehouck en Katrien Van Petegem verdienen een bijzondere vermelding voor de morele steun en inspiratie bij het plannen van experimenten of de verwerking ervan. Een aantal mensen staken ook letterlijk de handen uit de mouwen. Angelica Alcantara, Viki Vandomme en Robbe De Beelde van de TEREC en Martine Van Hove van het INBO hielpen me bij het versnijden en verpakken van gigantische hoeveelheden materiaal voor het multi-site experiment. Linda Stevens van het onthaal waakte over mijn berg dozen totdat DHL ze kwam opladen. Een speciale dank ook voor Yves Israel van het atelier die mijn gekste verzuchtingen kon inwilligen. 10 meter regenpijpen in stukjes van 5 cm versnijden? Geen probleem voor jou. Een systeem bedenken om plastiek gaas af te rollen en te versnijden? Een op maat gelast afrolsysteem was jouw creatieve en bijzonder effectieve oplossing voor mijn probleem. Verder zou ik ook graag Prof. Dr. Paul Simons van de faculteit Diergeneeskunde willen bedanken voor het bezorgen van allerlei eigenaardige attributen voor de experimenten, zoals schedels, hoeven en vacht van runderen en paarden. Ook de rondleiding in jullie gebouw was erg verhelderend (sinds ik weet welk hard werk studenten bij jullie moeten verrichten heb ik geen medelijden meer met biologie-studenten met natte voeten…), maar het werd me meteen ook duidelijk dat diergeneeskunde (en vooral het bloed dat daarbij komt kijken) niet meteen mijn ding is. Ook mocht ik ook de serres van het VLIZ gebruiken voor mijn serre- experimenten en kon ik op technische ondersteuning rekenen van Frank Broucke, Michiel T'Jampens en André Cattrijsse die ervoor zorgden dat mijn planten het overleefden. Het ANB liet me toe om in een aantal Vlaamse topreservaten experimenten te doen, vaak in het bijzijn van een legertje studenten. Een speciale vermelding voor Johan Lamaire, Koen Marechal en Karel Molenberghs, de boswachters van respectievelijk de Westhoek, het Zwin en de Kalmthoutse heide is dan ook op zijn plaats. Ook in dit kader zou ik Bram D'hondt speciaal willen bedanken. Ten eerste om ons als trotse conservator te laten experimenteren in 'jouw eigen' Natuurpunt-reservaat Doeveren, en ten tweede voor de medewerking in het allerlaatste experiment dat ik in het kader van mijn doctoraat verricht heb. Niet alleen was het voor mij bijzonder aangenaam om eindelijk nog eens buiten te kunnen komen na een lange periode van schrijven, ook was je een grote hulp bij het opstellen van het experiment en het vinden van een geschikte ii weide en mest. Nu van mest gesproken, voor wie het nog niet wist, mijn doctoraat handelt hoofdzakelijk over mest (wie een gigantische afkeer heeft voor fecale residues ligt dit werk dan best ook terug terzijde). In dat opzicht verdienen de vele dieren die me mest leverden of gewillig zaden van Geel zonneroosje naar binnen werkten een speciale vermelding. Het moet ergens middenin mijn doctoraat geweest zijn dat mijn promotors' subtiele hints over secundaire dispersie door mestkevers een ideetje deden kiemen. Dat ideetje groeide uiteindelijk uit tot een ALTER-Net multisite experiment met een financiering voor onderzoeksmateriaal. I would like to express my sincerest gratitude to all collaborators in this MSE-project! You all decided to trust me with such kind of project (although I realize that at that time I hadn't any dung beetle related publications, and even not that many as a first author) and donate much of your valuable time. Hopefully the resulting paper (chapter 8) will get published soon. Thank you Réka, Christophe, Tessa, Isabelle, Jörn, Constantin, Carsten, Amy, Boros, Marco, Pierre, Märt, Rosa, Enno, Geoffrey, Jörg, Jorge, Elham, Elena, Iraj, Dirk, Eleanor, László, Jutta, Pejman and Stefano! I really believe in such kinds of scientific cooperations, maybe we will meet again in a future project? Ik werd in 2011 door de Universiteit Gent aangeworven als assistent wat niet alleen inhield dat ik de kans kreeg om onderzoek te doen, maar ook ingeschakeld werd bij lesopdrachten. Dankzij een vorig leven bij het INBO wist ik al redelijk goed hoe aan dat onderzoek te beginnen, maar die lesopdrachten, dat was iets nieuw. Uiteindelijk bleek dat me wel te liggen, buiten het feit dan dat die studenten elk jaar jonger lijken te worden… Een speciale bedanking ook voor mijn collega-assistent Katrien Van Petegem voor de vele voorbereidingsweekendjes en -dagen voor de excursies. Een jaarlijks terugkerende constante was ook de organisatie van de stage. Een dikke merci ook voor de vele collega's die me het werk wat gemakkelijker maakten en zich vrij wilden maken tijdens het veldseizoen om onze studenten aan te steken met jullie enthousiasme! Gesproken over studenten, dit werk had er ook nooit in deze vorm gelegen zonder de waardevolle hulp van heel wat studenten. Tijdens mijn assistentschap had ik het geluk om met 5 bachelor- en 5 masterstudenten samen te werken in het kader van hun bachelor- en/of masterproef. Sanne Govaert, Febe Noppe, Manon Pien, Stijn Quidé en Benjamin Vercaempst deden zowat al het praktisch werk voor hoofdstuk 2 in het kader van hun iii bachelorproef. Harti Ningsih Suyoto studied the zoochorous dispersal potential of Helianthemum nummularium in her master thesis. The data of this thesis and supplementary growth experiments were united in chapter 6. Pieter-Jan D'hondt onderzocht de vegetatie-successie aan de Ijzermonding. Ook al past dit onderwerp niet echt in dit doctoraat, toch zou ik je graag willen bedanken voor je inzet bij het vele veldwerk dat je verricht hebt. Hopelijk vind ik ooit de tijd om iets meer met de resultaten te doen. Stefan Van Damme voerde in het kader van zijn master-thesis een piloot- experiment binnen het ALTER-Net multi-site experiment. Dankzij Stefan's input weten we welke methode werkt (en vooral ook wat niet werkt) en kwam er een duidelijk protocol voor de uitvoering van het MSE. Met dit gloednieuwe protocol konden Stijn Quidé en Thomas Van Loo dan weer aan de slag. Zij voerden het Belgische deel van de experimenten uit in 2 verschillende studiegebieden én hebben dat dan nog eens gerepliceerd over 2 seizoenen (zie onze impressionante paper in chapter 7!).
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