UNIVERSITÀ DEGLI STUDI DI TRIESTE XXXIII CICLO DEL DOTTORATO DI RICERCA IN AMBIENTE E VITA Global changes in harsh environments: effects on plant species and communities from a trait- analysis perspective. Settore scientifico-disciplinare: BIO/03 DOTTORANDO STEFANO VITTI COORDINATORE PROF. GIORGIO ALBERTI SUPERVISORE DI TESI DOTT. VALENTINO CASOLO CO-SUPERVISORE DI TESI DOTT. FRANCESCO BOSCUTTI ANNO ACCADEMICO 2019/2020 - 2 - STATEMENT OF ORIGINAL CONTRIBUTION This research thesis is an original contribution to the field of ecology and it is submitted as the final outcome of my Ph.D. project. It focused on linking plant responses to global climate changes to harsh environmental conditions, which determine remarkable species responses, from cell to community level, by means of physiological and morphological plant responses. I joined the Plant Biology Laboratory, Dipartimento di Scienze AgroAlimentari, Ambientali ed Animali (DI4A), University of Udine, Italy. My supervisor, Dr. Valentino Casolo, and co-supervisor Dr. Francesco Boscutti, always care for my work professionally, and their participation is revealed to be of fundamental relevance. This dissertation contains the papers, and other related works, I produced during my Ph.D. program. According to my academic discipline (BIO/03 - ENVIRONMENTAL AND APPLIED BOTANY), it is divided into main works (Chapter 1, 2, 3, 4) and other minor works (n.2), as follow: Chapter 1 Vitti, S., Pellegrini, E., Casolo, V., Trotta, G., & Boscutti, F. (2020). Contrasting responses of native and alien plant species to soil properties shed new light on the invasion of dune systems. Journal of Plant Ecology, rtaa052. doi: 10.1093/jpe/rtaa052; Chapter 2 Boscutti, F., Vitti, S., Casolo, V., Roppa, F., Tamburlin, D., & Sponza, S. (2019). Seagrass meadow cover and species composition drive the abundance of Eurasian wigeon (Mareca penelope L.) in a lagoon - 3 - ecosystem of the northern Adriatic Sea. Ecological Research, 34(2), 320– 327. doi: 10.1111/1440-1703.1070; Chapter 3 Population density modulates the response of morphological traits and non-structural carbohydrates in Vaccinum myrtillus when exposed to different rain regimes. Original draft, not submitted. Chapter 4 Trifilò, P., Kiorapostolou, N., Petruzzellis, F., Vitti, S., Petit, G., Lo Gullo, M.A., Nardini, A. and Casolo, V. (2019). Hydraulic recovery from xylem embolism in excised branches of twelve woody species: Relationships with parenchyma cells and non-structural carbohydrates. Plant Physiology and Biochemistry, 139, 513–520. doi: 10.1016/j.plaphy.2019.04.013. In addition to the above manuscripts, as previously said, other outputs have been presented at national - international conferences (for details see Appendix): • Vitti, S., Boscutti, F., Pellegrini, E., & Casolo, V. (2018). Study of plant traits, soil features and plant invasion in sandy beach of Grado and Marano lagoon (northern Adriatic Sea). 113° Congresso della Società Botanica Italiana - V International Plant Science Conference (IPSC). abstracts keynote lectures, communications, posters. Università di Salerno, Fisciano (SA). 978-88-85915-22-0. abstracts keynote lectures, communications, posters. Fisciano (SA): Università di Salerno. • Vitti, S., Boscutti, F., Casolo, V., & Sponza, S. (2019). Seagrass - waterbirds interactions in a lagoon ecosystem of the northern Adriatic Sea. 114° Congresso della Società Botanica Italiana - VI International Plant Science Conference (IPSC). abstracts keynote - 4 - lectures, communications, posters. Università di Padova, Padova. 978-88-85915-23-7. abstracts keynote lectures, communications, posters. Padova: Università di Padova; - 5 - ACKNOWLEDGEMENTS My most important thank goes to my supervisor, Dr. Valentino Casolo, and my co-supervisor, Dr. Francesco Boscutti, for being always helpful, professional, respectful, and full of wise advice, giving me a warm welcome to their research group. They taught me much about botany and ecology, also permitting me to discover fantastic unknown places during field activities. I thank the staff of Friuli Venezia Giulia Region for the administrative support, Dr. Glauco Vicario, and Dr. Stefano Sponza for field logistic support. I thank my office-mate, Antonio and Marco, particularly for recreational moments. They are really funny guys. Last but not least, I thank my family for morale and mental support, and thanks to Sara, the person who was most able to understand me in difficult moments. With her, I understand that, after all, the obstacles exist to be overcome. - 6 - - 7 - FOREWORD Climate is the average of the weather conditions at a particular point on the Earth, and it is expressed in terms of expected temperature, rainfall, and wind conditions based on historical observations (Riedy 2016). The Earth’s climate has always changed and the history of life on Earth is closely associated with environmental change on multiple spatial and temporal scales (Davis 2001). The success of human societies depends mainly on the living components of systems (Pecl et al. 2017). In the case of environmental conditions changes, for marine, freshwater, and terrestrial species, the first response is often a shift in location, to stay within ecologically optimal environmental conditions (Chen et al. 2011; Lenoir and Svenning 2015). At the cooler extremes of their distributions, species are moving poleward, whereas range limits are contracting at the warmer range edge, where temperatures are no longer tolerable. On land, species are also moving to cooler, higher elevations; in the ocean, they are moving to colder water at greater depths (Pecl et al. 2017). At a time when the world is anticipating unprecedented increases in human population growth and demands, the ability of natural ecosystems to deliver ecosystem services is being challenged by the largest climate-driven global redistribution of species, which is expected to influence climate feedbacks via changes in albedo, biologically driven sequestration of carbon from the atmosphere to the deep sea (the “biological pump”), and the release of greenhouse gases (Prentice et al. 2015). - 8 - Species are affected by climate in many ways, including range shifts, changes in relative abundance, and changes in activity timing and microhabitat use (Williams et al. 2008; Bates et al. 2014) and the geographic distribution of any species depends upon its environmental tolerance, dispersal constraints, and biological interactions with other species (Peterson et al. 2011). The race to save biodiversity is being lost, and it is being lost because the factors contributing to its degradation are more complex and powerful than those forces working to protect it (Wood et al. 2000). - 9 - TABLE OF CONTENT STATEMENT OF ORIGINAL CONTRIBUTION _________________________ - 3 - AKNOLEDGEMENTS _______________________________________________ - 6 - FOREWORD _______________________________________________________ - 8 - INTRODUCTION __________________________________________________ - 13 - Climate change and effects of plants in a changing precipitation scenario ________ - 13 - Biological invasion: invasion mechanisms in harsh environments _______________ - 15 - Effects of biological interactions ___________________________________________ - 17 - THESIS’ AIMS ____________________________________________________ - 19 - THESIS’S STRUCTURE ____________________________________________ - 20 - CHAPTER 1_______________________________________________________ - 23 - Contrasting responses of native and alien plant species to soil properties shed new light on the invasion of dune systems ___________________________________________ - 23 - CHAPTER 2_______________________________________________________ - 61 - Seagrass meadow cover and species composition drive the abundance of Eurasian wigeon (Mareca penelope L.) in a lagoon ecosystem of the northern Adriatic Sea ___ - 61 - CHAPTER 3_______________________________________________________ - 87 - Population density modulates the response of morphological traits and non-structural carbohydrates in Vaccinum myrtillus when exposed to different rain regimes _____ - 87 - CHAPTER 4______________________________________________________ - 120 - Hydraulic recovery from xylem embolism in excised branches of twelve woody species: relationships with parenchyma cells and non-structural carbohydrates _________ - 120 - CONCLUSIONS __________________________________________________ - 155 - APPENDIX ______________________________________________________ - 159 - Study of plant traits, soil features and plant invasion in sandy beach of Grado and Marano lagoon (Northern Adriatic Sea) ___________________________________ - 160 - Seagrass - waterbirds interactions in a lagoon ecosystem of the Northern Adriatic Sea_ - 164 - REFERENCES ___________________________________________________ - 168 - - 10 - “It is not the strongest of the species that survives, nor the most intelligent that survives. It is the one that is the most adaptable to change.” Charles Robert Darwin (12 February 1809 – 19 April 1882) INTRODUCTION Climate change and the effects of plants in a changing precipitation scenario Although it appeared on Earth relatively recently, man has affected most of the biosphere by means of its activities. Since the beginning of the industrial revolution, in England in the late 18th century, pollution due to human activities has been steadily increasing across the world. After the Second World War, there was an acceleration of this phenomenon: technological development has led