MEDITERRANEAN ODORSCAPES: Aphrodite N. Kantsa
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MEDITERRANEAN ODORSCAPES: THE ROLE OF PLANTS’ VOLATILE ORGANIC COMPOUNDS IN POLLINATION NETWORKS Aphrodite N. Kantsa Biologist, MSc Doctoral Dissertation submitted to the Department of Geography of the University of the Aegean defended in public on Thursday 29 September 2016 1 page intentionally left blank 2 ΜΕΣΟΓΕΙΑΚΑ ΟΣΜΟΤΟΠΙΑ: Ο ΡΟΛΟΣ ΤΩΝ ΑΡΩΜΑΤΩΝ ΤΩΝ ΦΥΤΩΝ ΣΤΑ ΔΙΚΤΥΑ ΕΠΙΚΟΝΙΑΣΗΣ Αφροδίτη Ν. Καντσά Βιολόγος, MSc Διδακτορική Διατριβή υποβληθείσα στο Τμήμα Γεωγραφίας του Πανεπιστημίου Αιγαίου Ημερομηνία δημόσιας υποστήριξης: Πέμπτη 29 Σεπτεμβρίου 2016 3 Kantsa, A. 2016. Mediterranean odorscapes: The role of plant volatile organic compounds in pollination networks. Doctoral Dissertation, University of the Aegean, Mytilene (pp. 225). Καντσά, Α. 2016. Μεσογειακά οσμοτοπία: Ο ρόλος των αρωμάτων των φυτών στα δίκτυα επικονίασης. Διδακτορική Διατριβή, Πανεπιστήμιο Αιγαίου, Μυτιλήνη (σελ. 225). Η έγκριση της διδακτορικής διατριβής από το Τμήμα Γεωγραφίας του Πανεπιστημίου Αιγαίου δεν υποδηλώνει την αποδοχή των γνωμών της συγγραφέως (Ν. 5343/1932, Άρθρο 202, Παράγραφος 2). 4 FUNDING This research has been co-financed by the EU (European Social Fund) and Greek national funds through the Operational Program “Education and Lifelong Learning” of the National Strategic Reference Framework – Research Funding Program: Heraclitus II (2324-1/WP17/30340). ΧΡΗΜΑΤΟΔΟΤΗΣΗ H παρούσα έρευνα έχει συγχρηματοδοτηθεί από την Ευρωπαϊκή Ένωση (Ευρωπαϊκό Κοινωνικό Ταμείο - ΕΚΤ) και από εθνικούς πόρους μέσω του Επιχειρησιακού Προγράμματος «Εκπαίδευση και Δια Βίου Μάθηση» του Εθνικού Στρατηγικού Πλαισίου Αναφοράς (ΕΣΠΑ) – Ερευνητικό Χρηματοδοτούμενο Έργο: Ηράκλειτος ΙΙ. 5 page intentionally left blank 6 THESIS COMMITTEE / ΕΠΙΤΡΟΠΗ ΔΙΑΤΡΙΒΗΣ Advisory Committee / Συμβουλευτική Επιτροπή 1. Theodora Petanidou, Professor, Department of Geography, University of the Aegean (Dissertation Advisor). 2. Themistokles Lekkas, Professor Emeritus, Department of Environment, University of the Aegean. 3. Jens M. Olesen, Professor, Department of Biosciences, Aarhus University. Examiners / Εξεταστές 1. Theodora Petanidou, Professor, Department of Geography, University of the Aegean (Dissertation Advisor). 2. Themistokles Lekkas, Professor Emeritus, Department of Environment, University of the Aegean. 3. Jens M. Olesen, Professor, Department of Biosciences, Aarhus University. 4. Robert A. Raguso, Professor, Department of Neurobiology and Behavior, Cornell University. 5. Stefanos P. Sgardelis, Professor, School of Biology, Aristotle University of Thessaloniki. 6. Panayiotis Dimitrakopoulos, Associate Professor, Department of Environment, University of the Aegean. 7. Thomas Tscheulin, Assistant Professor, Department of Geography, University of the Aegean. 7 PUBLICATIONS AND CONFERENCE CONTRIBUTIONS Kantsa A, et al. Floral sensory diversity and the structure of plant–pollinator networks (manuscript in prep). Kantsa A, et al. The insect–volatilome network in a Mediterranean community (manuscript in prep). Kantsa A, Raguso RA, Dyer AG, Olesen JM, Petanidou T. Smells like purple: community-wide integration of floral colour and scent (under review). Kantsa A, Sotiropoulou S, Vaitis M, Petanidou T. 2015. Plant volatilome in Greece: a review on the properties, prospects, and chemogeography. Chemistry & Biodiversity, 12(10): 1466-1480. Kantsa A, Raguso RA, Olesen JM, Kalantzi O-I, Lekkas T, Petanidou T. 2014. Ο λειτουργικός ρόλος των αρωμάτων των φυτών στα επικονιαστικά δίκτυα (The functional role of plant scents in pollination networks) (oral presentation). Proceedings of the 7th Conference of the Hellenic Ecological Society, Mytilene, Greece, 9-12 October, p. 97 (in Greek). Kantsa A, Raguso RA, Olesen JM, Petanidou T. 2014. Plant volatiles and floral visitors in a Mediterranean community: A network approach (oral presentation). Gordon Research Seminar on Floral and Vegetative Volatiles. Ventura, CA, USA, 25-26 January. Kantsa A, Vaitis M, Petanidou T. 2010. AROMAbase: Η πρώτη βάση δεδομένων για τη χημική ποικιλότητα των αρωμάτων της ελληνικής χλωρίδας (AROMAbase: The first database for the chemical diversity of scents of the Greek flora) (poster). Proceedings of the 5th Conference of the Hellenic Ecological Society, Patras, Greece, 7-10 October, p. 152 (in Greek). Kantsa A, Petanidou T, Raguso RA. 2009. Exploring floral scents and their role in structuring plant– pollinator interaction networks (poster). Gordon Research Conference on Floral and Vegetative Volatiles. Oxford, UK, 9-14 August. 8 Στον πατέρα μου. […] Γύρω απ’ την ολοπράσινη επιτυχία των φύλλων Οι πεταλούδες ζουν μεγάλες περιπέτειες Ενώ η αθωότητα Ξεντύνεται το τελευταίο της ψέμα Γλυκιά περιπέτεια Γλυκιά Η Ζωή. Οδυσσέας Ελύτης, Δεύτερη Φύση, ΙΙ 9 page intentionally left blank 10 ACKNOWLEDGEMENTS “…a very little thief of occasion will rob you of a great deal of patience…” A well-respected scientist once said to me, in a joking manner, that you spend all these years doing research in order to finally write a prologue. Back then I laughed, but I’m afraid that I now have to acknowledge the overwhelming satisfaction of this very moment. Another well-respected scientist I know, stoically (or perhaps Socratically) concluded that although when you get your first degree you think that you know everything, when you get your PhD, you confirm that you actually know nothing. Being fully conscious of these two conditions, I feel like thanking all those people who patiently helped me through this intellectual achievement . It all started with Prof. Theodora Petanidou. She first introduced me to the floral sensory ecology by suggesting measuring the odorscape created by the insect-pollinated flowers in an entire community, and study its effects on the local pollination network. I was easily fascinated by the idea. I would have probably thought about it a bit more, if I knew the amount of work required, nevertheless my decision would not have been any different. If I ever become a decent ecologist, it will be largely due to her ‘demonic’ instinct and profound knowledge on the Mediterranean nature, to the tough fieldwork training (no joke), to the endless conversations and the arguments we’ve had (hardcore as well), and to the education she’s constantly providing. She is probably the most cultured person I’ve ever met, and although I often feel perplexed by this, she is a precious mentor and friend. Professor Jens Olesen (Aarhus University) opened the gate to network analysis for me, being also a member of my Advisory Committee. My stay in Aarhus University in 2013 was one of the best memories I have from my PhD, and I deeply thank him for his hospitality, is trust, his sense of humor and his Scandinavian serenity that has been of help more than once. I owe warm thanks to Prof. Themistokles Lekkas (University of the Aegean), also a member of my Advisory Committee, who was among the first people who believed in this project, and made possible for me to use the infrastructure of the Laboratory of Water and Air Quality at the Department of Environment. Without his help, my hands-on training in GC/MS analysis would be unfeasible in Mytilene. Professor Robert Raguso (Cornell University) was generous and patient to provide detailed guidelines for scent collection and analysis, as well as moral support. Even from before we came into contact, I had been fascinated by the fact that he stood up for community-wide studies of floral scent and the integration of sensory data in pollination network analysis. Robert Raguso has taught me a lot and I profoundly thank him for this. Professor Stefanos Sgardelis (Aristotle University of Thessaloniki) was a member of the Examination Committee as well as a great advisor regarding statistical analysis. I thank him for the solutions and the ideas he offered and for tolerating my (mostly naïve) questions. Assistant Professor Thomas Tscheulin (University of the Aegean) has been a valuable friend since the very beginning of this journey. I thank him for being there, for listening to my ideas and troubles, for the advice and help he provided, for the fun and games at his Christmas dinners, and for his questions and comments regarding this thesis as a member of the Examination Committee. I thank Assoc. Prof. Panagiotis Dimitrakopoulos (University of the Aegean) for accepting to participate in the Examination Committee, for his encouragement and his comments. 11 Associate Professor Adrian Dyer (Royal Melbourne Institute of Technology) undertook the colorimetric analyses of my dataset. I warmly thank him for the valuable lessons on visual ecology and insects’ sensory systems, as well as for his patience. Assistant Professor Michalis Vaitis (University of the Aegean) designed the geodatabase of the volatile chemodiversity of the plants in Greece and helped me with databasing my insect specimens. Professor Peter Davies (Cornell University) was the first one who showed to me in person how to process and analyze my scent specimens in the GC/MS lab. I am deeply thankful that he was next to me at the illuminating moment when the first peak, β-pinene, appeared on the monitor. I am grateful to Prof. Ante Vujić (University of Novi Sad) who identified the Syrphidae specimens, to Jelle Devalez and Marino Quaranta for identifying Hymenoptera, to Jos Dils for identifying Bombyliidae, and to Assis. Prof. Thomas Tscheulin for identifying several Coleoptera specimens. Dr. Anastasia Stefanaki and Katerina Goula confirmed the identification of several plant specimens. Associate Professor Thanassis Kallimanis (Aristotle University