Effects of artificial light at night (ALAN) on interactions between aquatic and terrestrial ecosystems Alessandro Manfrin Freie Universität Berlin Berlin, 2017 Effects of artificial light at night (ALAN) on interactions between aquatic and terrestrial ecosystems Inaugural-Dissertation to obtain the academic degree Doctor of Philosophy (Ph.D.) in River Science submitted to the Department of Biology, Chemistry and Pharmacy of Freie Universität Berlin by ALESSANDRO MANFRIN from Rome, Italy Berlin, 2017 I This thesis work was conducted during the period 29th September 2013 – 21st February 2017, under the supervision of PD. Dr. Franz Hölker (Leibniz-Institute of Freshwater Ecology and Inland Fisheries Berlin), Dr. Michael T. Monaghan (Leibniz- Institute of Freshwater Ecology and Inland Fisheries Berlin), Prof. Dr. Klement Tockner (Freie Universität Berlin and Leibniz-Institute of Freshwater Ecology and Inland Fisheries Berlin), Dr. Cristina Bruno (Edmund Mach Foundation San Michele all´Adige) and Prof. Dr. Geraldene Wharton (Queen Mary University of London). This thesis work was conducted at Freie Universität Berlin, Queen Mary University of London and University of Trento. Partner institutes were Leibniz-Institute of Freshwater Ecology and Inland Fisheries of Berlin and Edmund Mach Foundation of San Michele all´Adige. 1st Reviewer: PD. Dr. Franz Hölker 2nd Reviewer: Prof. Dr. Klement Tockner Date of defence: 22nd May 2017 II The SMART Joint Doctorate Programme Research for this thesis was conducted with the support of the Erasmus Mundus Programme1, within the framework of the Erasmus Mundus Joint Doctorate (EMJD) SMART (Science for MAnagement of Rivers and their Tidal systems). EMJDs aim to foster cooperation between higher education institutions and academic staff in Europe and third countries with a view to creating centres of excellence and providing a highly skilled 21st century workforce enabled to lead social, cultural and economic developments. All EMJDs involve mandatory mobility between the universities in the consortia and lead to the award of recognised joint, double or multiple degrees. The SMART programme represents a collaboration among the University of Trento, Queen Mary University of London, and Freie Universität Berlin. Each doctoral candidate within the SMART programme has conformed to the following during their 3 years of study: (i) Supervision by a minimum of two supervisors in two institutions (their primary and secondary institutions). (ii) Study for a minimum period of 6 months at their secondary institution (iii) Successful completion of a minimum of 30 ECTS of taught courses (iv) Collaboration with an associate partner to develop a particular component / application of their research that is of mutual interest. (v) Submission of a thesis within 3 years of commencing the programme. 1This project has been funded with support from the European Commission. This publication reflects the views only of the author, and the Commission cannot be held responsible for any use which may be made of the information contained therein III Acknowledgements Acknowledgements First I want to thank The SMART supervisory board: Prof. Klement Tockner, Prof. Angela Gurnell and Prof. Guido Zolezzi, for giving me the unique opportunity to take part in the SMART doctoral program. I do not have words to thank my supervisors Dr. Michael T. Monaghan, PD. Dr. Franz Hölker, Prof. Klement Tockner, Dr. Cristina Bruno and Prof. Geraldene Wharton for being a source of inspiration during these three years. Very special thanks go to the three best colleagues and friends I have ever worked with: Dr. Stefano Larsen, Dr. Roy van Grunsven and (soon Dr.) Maja Grubišić. I am grateful to the colleagues, students and trainees at the IGB who helped me in the field, in the laboratory, or with fruitful discussions. Particularly I want to thank: Ann- Christin Honnen, Valentyna Inshyna, Viktor Baranov, Babette Pohlmann, Nina-Sophie Weiss, Nadine Weiss, Liliana Lehmann, and many others from the Monday journal/discussion group. Special thanks go to Thomas Mehner, Kate Laskowski, Gabriel Singer, Kirsten Pohlmann and Francesca Pilotto for their help with statistics and for their, necessarily critical, discussions on my research. Special thanks go to Elizabeth Perkin and Abel Machado for their important comments on the manuscripts and to Lisa Angermann for helping me with the german translation of the summary. I also want to thank Stefan Heller, Lorenzo Forti and Martino Salvaro for their help in setting up of the experiments; and Sibylle Schroer, Anika Brüning, Tobias Degen, Stephanie Holzhauer and the entire “Verlust der Nacht” group for such incredible adventure. I want to say thank to Stefanie Burkert, Frau Katrin Lhemann, Frau Marlis Lange and Marina Rogato for their precious support on administrative issues. The colleagues and the other PhD students from IGB and the SMART program. Especially I want to thank Çağrı Gökdemir, Alex Lumsdon, Pascal Bodmer and Oleksandra Shumilova with whom I particularly shared this unique experience. I want to thank my beloved friends Massimo, Simone, Valeria, Alessio, Ruby, Manolo, Fabrizio, Tommy, Massi, Rene, Fanny, Patty, Carolina, Arianna, Annika, Loredana, Cristina and Benito for their continuous support. Finally, and most of all, I want to thank my family Fiorella, Emilio and Millie for constantly encouraging me and being always present when I needed. IV “Research will work for you, but finding cheap flights is still an important skill in science, too!” M. T. M. V Table of contents Table of contents Table of contents...................................................................................................... 1 Summary .................................................................................................................. 4 Zusammenfassung .................................................................................................. 7 Thesis outline and collaboration statement ....................................................... 11 1. General introduction .......................................................................................... 13 1.1. Artificial light at night (ALAN) ......................................................................... 13 1.2. Effect of ALAN on organisms ........................................................................ 15 1.3. Effect of ALAN on the coupled aquatic-terrestrial ecosystems ....................... 16 1.4 Knowledge gaps.............................................................................................. 18 1.5 Thesis aims and approach .............................................................................. 21 1.6 References ...................................................................................................... 24 2. Artificial light at night (ALAN) affects structural and functional aspects of macroinvertebrate assemblages: a field experiment in a previously ALAN- naïve area ............................................................................................................... 31 2.1 Abstract ........................................................................................................... 32 2.2 Introduction ..................................................................................................... 33 2.3 Methods .......................................................................................................... 34 2.3.1 Study site .................................................................................................. 34 2.3.2 Animal collection and experimental design ............................................... 36 2.3.3 Feeding groups ......................................................................................... 37 2.3.4 Data analysis design ................................................................................. 38 2.3.5 Statistical analysis .................................................................................... 38 2.4 Results ............................................................................................................ 40 2.4.1 ALAN-naïve communities ......................................................................... 40 2.4.2 ALAN-exposed communities .................................................................... 41 2.4.3 Community resilience post-ALAN ............................................................. 41 2.5 Discussion ....................................................................................................... 45 2.5.1 ALAN-naïve communities ......................................................................... 45 2.5.2 ALAN-exposed communities and community resilience post-ALAN ......... 47 2.5.3 Seasonality ............................................................................................... 47 2.5.4 Ecological implications of ALAN in freshwater ecosystems ...................... 49 2.6 Acknowledgements ......................................................................................... 50 2.7 References ...................................................................................................... 51 1 Table of contents 3. Artificial light at night alters flux across ecosystem boundaries and community structure in the recipient ecosystem ............................................... 57 3.1 Abstract ........................................................................................................... 58 3.2 Introduction ....................................................................................................