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Università degli Studi di Cagliari DOTTORATO DI RICERCA BOTANICA AMBIENTALE E APPLICATA Ciclo XXVI Germination niche of Sardinian endemic species in mountain riparian deciduous forests BIO/03 Presentata da: Dr. Marco Porceddu Coordinatore Dottorato Prof. Gianluigi Bacchetta Tutor Prof. Gianluigi Bacchetta Prof. Hugh W. Pritchard PhD. Efisio Mattana Esame finale anno accademico 2012 – 2013 A sa famíglia e sa isposa mia cun tottu su coru 3 La presente tesi è stata prodotta durante la frequenza del corso di dottorato in BOTANICA AMBIENTALE E APPLICATA dell’Università degli Studi di Cagliari, a.a. 2012/2013 - XXVI ciclo, con il supporto di una borsa di studio finanziata con le risorse del P.O.R. SARDEGNA F.S.E. 2007-2013 - Obiettivo competitività regionale e occupazione, Asse IV Capitale umano, Linea di Attività l.3.1 “Finanziamento di corsi di dottorato finalizzati alla formazione di capitale umano altamente specializzato, in particolare per i settori dell’ICT, delle nanotecnologie e delle biotecnologie, dell'energia e dello sviluppo sostenibile, dell'agroalimentare e dei materiali tradizionali”. Marco Porceddu gratefully acknowledges Sardinia Regional Government for the financial support of her PhD scholarship (P.O.R. Sardegna F.S.E. Operational Programme of the Autonomous Region of Sardinia, European Social Fund 2007-2013 - Axis IV Human Resources, Objective l.3, Line of Activity l.3.1.)”. 4 Index Summary ......................................................................................................................................... 9 General Introduction and Literature Review .................................................................................. 11 Seed structure ............................................................................................................................ 11 Seed germination ....................................................................................................................... 13 Seed dormancy .......................................................................................................................... 15 Evolutionary trends of seeds structure and dormancy ................................................................. 18 Thermal time model ................................................................................................................... 20 Mediterranean climate ............................................................................................................... 22 Mediterranean basin................................................................................................................... 23 Climate change .......................................................................................................................... 25 Aims .......................................................................................................................................... 27 References ................................................................................................................................. 28 Chapter I - Thermal niche for in situ seed germination by Mediterranean mountain streams: model prediction and validation for Rhamnus persicifolia seeds1 .............................................................. 32 Abstract ..................................................................................................................................... 32 Introduction ............................................................................................................................... 33 Material and Methods ................................................................................................................ 36 Study species.......................................................................................................................... 36 Seed lot details ....................................................................................................................... 36 Germination tests under controlled conditions ....................................................................... 38 Field experiments .................................................................................................................. 38 Data analysis ......................................................................................................................... 39 Results ....................................................................................................................................... 41 Seed germination under controlled conditions ....................................................................... 41 Thermal requirement for germination .................................................................................... 42 Seed germination in the field ................................................................................................. 45 Soil heat sum and thermal niche for in situ seed germination ................................................. 47 Discussion ................................................................................................................................. 49 Type of dormancy .................................................................................................................. 49 Thermal requirements for germination .................................................................................. 50 Soil heat sum and thermal niche for in situ seed germination ................................................. 51 Conclusions ............................................................................................................................... 53 Chapter II - Multiphasic thermal parameters for embryo growth, seed dormancy loss and germination in Aquilegia barbaricina ................................................................................................................ 60 Abstract ..................................................................................................................................... 60 Introduction ............................................................................................................................... 61 Materials and Methods .............................................................................................................. 62 Study species ......................................................................................................................... 62 Seed lot details ...................................................................................................................... 63 Germination tests .................................................................................................................. 63 Embryo measurements ........................................................................................................... 64 Thermal time analyses ........................................................................................................... 65 Statistical analysis ................................................................................................................. 67 Results ...................................................................................................................................... 67 Embryo growth, endosperm rupture and seed germination .................................................... 67 Thermal time approach on embryo growth ............................................................................ 71 Thermal time approach on seed germination ......................................................................... 73 Discussion ................................................................................................................................. 76 Type of dormancy .................................................................................................................. 76 Multiphasic seed germination ................................................................................................ 77 6 Thermal thresholds for embryo growth and seed germination ................................................ 78 Conclusions ............................................................................................................................... 80 Chapter III - Sequential temperature control of multiphasic growth and germination of Paeonia corsica seeds.............................................................................................................................................. 86 Abstract ..................................................................................................................................... 86 Study species and seedlot details ............................................................................................ 89 Experimental trials ................................................................................................................ 89 Embryo measurements ........................................................................................................... 90 Endosperm rupture and radicle emergence ............................................................................ 91 Epicotyl dormancy release ..................................................................................................... 91 Statistical analysis ................................................................................................................. 91 Results ......................................................................................................................................