Spatial urban and peri-urban biodiversity patterns Olga Tzortzakaki PhD Thesis Department of Biology University of Patras 2018 2 Χωρικά πρό τυπά άστική ς κάι περιάστική ς βιόπόικιλό τήτάς Όλγά Τζωρτζάκάκή Διδάκτόρική Διάτριβή Τμήμά Βιόλόγίάς Πάνεπιστήμιό Πάτρών 2018 3 4 Examining Committee Sinos Giokas, Associate Professor, University of Patras (Supervisor) Vassiliki Kati, Associate Professor, University of Ioannina (Advisory Committee) Konstantinos Nikolakopoulos, Associate Professor, University of Patras (Advisory Committee) Maria Panitsa, Assistant Professor, University of Patras Savvas Kazantzidis, Senior Researcher, Hellenic Agricultural Organization “DEMETER” Robert Nudds, Senior Lecturer, Faculty of Biology, Medicine & Health, University of Manchester, UK Benoît Fontaine, Researcher, Muséum National d'Histoire Naturelle, France Επταμελής Εξεταστική Επιτροπή Σίνος Γκιώκας, Αναπληρωτής Καθηγητής, Πανεπιστήμιο Πατρών (Επιβλέπων) Βασιλική Κατή, Αναπληρώτρια Καθηγήτρια, Πανεπιστήμιο Ιωαννίνων (Τριμελής Συμβουλευτική Επιτροπή) Κωνσταντίνος Νικολακόπουλος, Αναπληρωτής Καθηγητής, Πανεπιστήμιο Πατρών (Τριμελής Συμβουλευτική Επιτροπή) Μαρία Πανίτσα, Επίκουρη Καθηγήτρια, Πανεπιστήμιο Πατρών Σάββας Καζαντζίδης, Τακτικός Ερευνητής, Ελληνικός Γεωργικός Οργανισμός «ΔΗΜΗΤΡΑ» Robert Nudds, Επίκουρος Καθηγητής, Τμήμα Βιολογίας, Ιατρικής και Υγείας, Πανεπιστήμιο Manchester, ΗΒ Benoît Fontaine, Ερευνητής, Εθνικό Μουσείο Φυσικής Ιστορίας, Γαλλία Approval of a dissertation for the obtainment of a PhD degree from the Department of Biology of the University of Patras does not imply acceptance of the author’s opinions. L. 5343/1932, article 202. Η έγκριση της διατριβής για την απόκτηση Διδακτορικού Διπλώματος από το Τμήμα Βιολογίας του Πανεπιστημίου Πατρών δεν υποδηλώνει την αποδοχή των γνωμών του συγγραφέα. Ν. 5343/1932, άρθρο 202. 5 6 Table of contents Acknowledgements .................................................................................................................. 12 Chapter 1. General introduction .......................................................................................... 15 1.1 The urban environment ............................................................................................. 16 1.2 Impacts of urbanisation on biodiversity .................................................................... 17 1.3 Species responses to urbanisation ............................................................................. 19 1.4 Biodiversity conservation in the urban environment ................................................ 20 1.5 Scope of the study ..................................................................................................... 20 1.6 Thesis outline ............................................................................................................ 22 References ............................................................................................................................ 23 Chapter 2. Study area........................................................................................................... 31 2.1 Study area .................................................................................................................. 32 2.2 The urbanisation gradient .......................................................................................... 34 References ............................................................................................................................ 37 Chapter 3. Seasonal patterns of urban bird diversity in a Mediterranean coastal city: the positive role of open green spaces ........................................................................................... 39 3.1 Introduction ............................................................................................................... 40 3.2 Methods ..................................................................................................................... 42 3.2.1 Study area and site selection .............................................................................. 42 3.2.2 Bird surveys ....................................................................................................... 43 3.2.3 Land-cover types ................................................................................................ 44 3.2.4 Data analysis ...................................................................................................... 44 3.3 Results ....................................................................................................................... 45 3.3.1. The urbanisation gradient .................................................................................. 45 3.3.2. Bird diversity ..................................................................................................... 46 3.3.3. Effects of land-cover types on bird diversity ..................................................... 50 3.3.4. Bird community structure .................................................................................. 51 3.4 Discussion ................................................................................................................. 53 3.4.1. The urban environment ...................................................................................... 53 3.4.2. Seasonal bird diversity ....................................................................................... 54 3.4.3. Effect of land-cover types on urban bird diversity ............................................ 54 3.4.4. Management implications .................................................................................. 56 References ............................................................................................................................ 58 7 Chapter 4. Butterfly diversity along the urbanisation gradient in a densely-built Mediterranean city: Land cover is more decisive than resources in structuring communities 65 4.1 Introduction ............................................................................................................... 66 4.2 Methods ..................................................................................................................... 67 4.2.1 Study area and sampling sites ............................................................................ 67 4.2.2 Butterfly surveys ................................................................................................ 69 4.2.3 Land cover and local habitat characteristics ...................................................... 69 4.2.4 Data analysis ...................................................................................................... 70 4.3 Results ....................................................................................................................... 71 4.3.1 Vegetation and habitat features along the urbanisation gradient ....................... 72 4.3.2 Butterfly diversity along the urbanisation gradient ........................................... 73 4.3.3 Drivers of butterfly diversity patterns ................................................................ 74 4.4 Discussion ................................................................................................................. 77 4.5 Conclusions ............................................................................................................... 80 References ............................................................................................................................ 81 Chapter 5: Bat diversity patterns along the urbanisation gradient ........................................... 87 5.1 Introduction ............................................................................................................... 88 5.2 Methods ..................................................................................................................... 89 5.2.1 Study area........................................................................................................... 89 5.2.2 Bat surveys and sound data analysis .................................................................. 91 5.2.3 Environmental variables .................................................................................... 93 5.2.4 Data analysis ...................................................................................................... 94 5.3 Results ....................................................................................................................... 95 5.3.1 The urban environment ...................................................................................... 95 5.3.2 Species identification and occurrence ................................................................ 96 5.3.3 Bat activity along the urbanisation gradient ...................................................... 98 5.3.4 Relationship between bat activity and environmental variables ...................... 100 5.4 Discussion ............................................................................................................... 103 References .......................................................................................................................... 108 Chapter 6: General discussion ............................................................................................... 115 6.1 Diversity along the urbanisation gradient ............................................................... 116 6.2 The effect of the environment on diversity